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Full text of "North American cestodes of the order Pseudophyllidea parasitic in marine and fresh water fishes"

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in 2013 



http://archive.org/details/northamericancesOOcoop_0 



NORTH AMERICAN CESTODES OF THE ORDER PSEUDOPHYLLIDEA 
PARASITIC IN MARINE AND FRESH WATER FISHES 



ARTHUR REUBEN COOPER 

A. B. University of Toronto, 1910 
A. M. University of Toronto, 1911 



THESIS 

Submitted in Partial Fulfillment of the Requirements for the 

Degree of 
DOCTOR OF PHILOSOPHY 
IN ZOOLOGY 

IN 

THE GRADUATE SCHOOL 

OF THE 

UNIVERSITY OF ILLINOIS 
1917 



C7^ 



UNIVERSITY OF ILLINOIS 
THE GRADUATE SCHOOL 



M 1.7 IQI 7 



I HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER MY SUPER- 
VISION by Arthur Reuben Coop er _ 

entitled lI.oxtk..A.m.erl.ca.n„ 

phyllide a Parasitic i n Marine a n d Fresh Water Fishes • „._ 

BE ACCEPTED AS FULFILLING THIS PART OF THE REQUIREMENTS FOR THE 

degree of Doctor of Philosophy _ 

c3l 




Recommendation concurred in :* 

LzZZZ. (IL^Ad^-....... 




Committee 



Final Examination* 



^Required for doctor's degree but not for master's. 

376654 



INDEX OF CONTENTS. 

Introduction 1. 

Historical Data 4* 

Explanation of Terms 8. 

Order PSEUDOPHYLL I DE A 9. 

1 Family, DIPHYLLOBOTHRI I DAE 12. 

1 Subfamily, LIGULINAE 16. 

1 Genus, Ligula 17. 

Ligula intestinalis (Linnaeus) 19. 

2 Genus, Schistocephalus 45. 

Schistocephalus solidus (0. F. Mueller) 45. 

2 Subfamily, HAPLOBOTHRI INAE subfam. nov 68. 

1 Genus, Haplobothrium 72. 

Haplobothrium globulif orme Cooper 73. 

3 Subfamily, CYATHOCSPHALINAE 90. 

1 Genus, Cyathocephalus 90. 

Cyathocephalus americanus sp.nov 91. 

4 Subfamily, MARS I POLE TR INAE subfam. nov 109. 

1 Genus, Marsipometra gen. nov 110. 

Mars i pome tra hastata (Linton) 112. 

5 Subfamily, TRIAENOPHORINAE 130. 

1 Genus, Triaenophorus 130. 

Triaenophorus sp.larv 133. 

2 Family, PTYCHOBOTHRI IDAE 140. 

1 Subfamily, PTYCHOBOTHRI INAE 143. 



1 Genus, Bothriocephalus s.str 145 

1 Species, B. scorpii (Mueller) 146 

2 n , B. claviceps (Goeze) 178 

3 " , B. cuspidatus sp.nov 188 

4 ,r , B. manubrif oralis (Linton) 193 

5 n , B. occidentalis (Linton) 205 

2 Genus, Clestobothrium 208 

Clestobothrium crassiceps (Rudolphi) 209 

2 Subfamily, AMPHICOTYLIHAE 244 

1 Genus, Abothrium 244 

1 Species, Abothrium ruffosum (Eatsch) 246 

2 Species, Abothrium crassum (Eloch) 264 

Explanation of Plates. 

Plates I-X. 

Vita* 



1 

INTRODUCTION 

Soon after commencing the study of Haplobothrium globuli- 
fjorme Cooper (1914 a and b) the writer saw that, apart from the 
early and somewhat brief reports and descriptions by Leidy and the 
later, but yet pioneer work of Linton on both marine and fresh- 
water species, very little had been done on the members of the or- 
der in America. Consequently the desire for an opportunity to 
work up other 3pecie3 which had in the meantime been collected at 
the Canadian Lake Biological Station on GeorgianjBay, located at 
Go-Home-Bay, Muskoka District, Ontario, and at the Marine Biologi- 
cal Station at St. Andrews, New Brunswick, grew with the feeling 
that something of a comprehensive nature ought to be undertaken in 
order not only to ascertain to what extent European species are to 
be found in thi3 continent but also to properly locate in the 
classification at least 3ome of the new forms formerly described 
especially by Linton. Altho the material then at hand was inves- 
tigated to a certain extent at the University of Toronto, it was 
not until the writer came to the University of Illinois that it was 
studied at all thoroughly with the aid of other material from the 
collection of the University of Illinois, under the care of 
Professor Henry 3. Ward, for comparison. 

Supplementary material, which in many cases was all that 
was available, was obtained by Dr. Ward from the United States 
National Museum and the Bureau of Animal Industry, but apart from 
a few vials no European specimens could be procured owing to the 
present international conflict. On account of the lack of the 
latter most of the determinations have been made with the aid of 



3 

the literature only, a fact which the writer feels may necessitate 
future changes in connnection with a few species which have been 
more or less tentatively regarded to be the same a3 those in Europe. 
In all cr.ses, however, the specific details of the American forms 
have been emphasized 30 that if changes have to be made later, the 

UKt\x> 

b^-sis for such will be at hand. The writer wculd N point out in 
this connection the comparative lack from a systematic standpoint 
of adequate descriptions of many of the European species which 
have been known for many years. It was thi3 fact which in the 
absence of the original material for comparison made the present 
work one attended with not a little difficulty. 

In the main the classification of the order adopted by the 
writer is that proposed by Luehe in 1902 in his "Revision meine3 
3othriocephaliden3ystemes" and later (1910) retained in "Die Su3s- 
wa33erfauna Deutschlands" with only a few modifications. The 
family of the Caryophyllaeidae is, however, not included, 30 that 
the order is considered to be rather that of Carus (1363), with, 
of course, Luehe' s later conceptions of the other families. One 
of the latter ha3 now again got to be modified considerably owing 
to the present 3tudy of two quite aberrant species, viz., Hap lob 0- 
thriurn fflobu li forme Cooper and Mar3ipometra ha-stata (Linton) which 
have been found by the writer to be very disturbing to the classi- 
fication, the former, in fact, possessing a scalex which bears 
strong resemblance to that of the order Trypauorhyncha . 

The writer wishes to here tender his thanks in the first 
place to the Biological Board of Canada for placing means and 
facilities at his disposal in connection with his earlier collect- 
ing at the above mentioned Canadian Biological Stations; to the 



3 

University of Illinois for the opportunity of collecting further 
material at the Marine Biological Laboratory at Woods Hole, Massa- 
chusetts, and at the Harpswell Laboratory, South Harpswell, Maine, 

during the summer of 1916, and to the staffs of these institutions 

r 

as well as to that of the Marine Laboratory of the United States 
Bureau of Fisheries at Woods Hole for a33i3tance and direction in 
connection with the same; to the Smithsonian Institute and the 
Bureau of Animal Industry, from whom valuable material was obtained 
for comparison, in the latter case thru the kind offices of 
Dr. C. W. Slies of the Hygienic Laboratory, Washington; and to the 
following investigators for alcoholic specimens: Professor 0. 
Fuhrman* University of Neuchatel, Professor Edwin Linton, Washing- 
ton and Jefferson College, Professor E. M. Walker, University of 
Toronto, Dr. H. J. Van Cleave, University of Illinois, Mr. H. R. 
Hill and Mr. R. P. Wodehouse. 

Finally to Professor H. B. Ward the v/riter wishes to ex- 
press his sincere indebtedness not only for the U3e of his exten- 
sive private library and collections and for his procuring rare 
books and specimens, but for his constant and stimulative interest 
in and valuable criticism of the work which has resulted in the 
following paper. 



I 

4 

HISTORICAL DATA 

Apart from Omeiin'3 (1790) collecting together the data- 
given by the older writers such as Linnaeus, Pallas, Mueller, 
Goeze, Bloch, Fabricius, Batsch, Schrank, and Abildgaard and 
Zeder's (1830 and 1803) treatises, the first mo3t important work 
on the bothriocephaiid cestodes was the "Entozoorum Historia 
Naturalis" by Rudolphi (1803-1810). In this he reviewed the 
earlier literature, making valuable comments on the same, and 
described species of Ligu la, Tria enophorus and 3othrio cephalu3 , 
the latter name being used for the first time. While Lamarck 
(1813) dealt with only the more common species, Rudolphi in hi 3 
second work of major importance, the "Entozoorum Synopsis" (1319), 
made some corrections of his earlier publication and further con- 
tributions in the way of a few new species. LeiLckart (1819), who 
did not receive Rudolphi '3 "Entozoorum" until after hi3 work was 
in print, dealt only with 3pecies of the genus Bothrio cephalus as 
conceived by Rudolphi, which then contained members not only of the 
Pseudophyllidea but also of the Tetraphyllidea and the Trypanorhyn- 
cha. Nitzsch (1834) briefly defined the species of the same genus, 
while later in the same publication Creplin (1839) dealt with them 
more in detail and erected the new genus Schistocephalus . Drummond 
(1838) was one of the first to report bothriocephalids from the 
British Isle3, while Bellingham (1844) and Thompson (1844) made 
further contributions , all three dealing with forms from Ireland. 
Eschricht (1841) published some of the earliest data on the internal 
anatomy of the group, and TCceliiker (1843) made a study of the 
development of the eggs of a few species. The next and perhaps 



o 

moat important work was that by Dujardin (1845) who, while follow- 
ing Rudolphi in the main, made many valuable additions from origin- 
al observations. Van Beneden (1849 and 1850) first essayed to 
erect a more comprehensive classification than had hitherto been 
used, and Die sing (1850) went much farther in his Subtribe I, 
Gymno/cothria, of Tribe IV, Bothriocephalidea , of Suborder I, 
Aprocta, of Order IV Cephalocotylea . Baird (1853) reverted to 
Rudolphi ' s brief system in listing forms from the British Museum. 
lYagener in two papsrs (1854 and 1857) gave studies on the develop- 
ment which even today are models of careful work and excellent 
illustrating. Leidy (1855 and 1856) was the first to report forms 
from America, while Weinland (1858) in his "E3say" made a few 
references to bothriocephaiids . Then, until Diesing (18S3) revised 
hi 3 classification nothing of systematic importance was published. 
01s3on (1867) was one of the first to report species from the 
Scandinavian countries; later (1876 and 1893) he made further con- 
tributions from the same source. After Trillemoes-Suhm' s (186S) 
studies on the development of Sen, dimorphus , came Duchamp's (1876) 
and Donnadieu's (1877) classical experiments on the life- hi stories 
of the Ligutes. In his Compendium Linatow (1878) brought together 
in a list the forms known up to that time. A few years later 
Fraipont (1880, 1881) published studies on the excretory system of 
a number of species which even to-day are perhaps the most impor- 
tant contributions in that direction. The nerv ous system was made 
the object of special inquiry by Lang (1881), while later it was 
dealt with more at length by Niemiec (1888) and Cohn (1898). After 
a period in which such works as Moniez's (1881) "Memoire", 
Z&chokke's (1884) researches, and Schaumsland 1 s (1885) studies of 



6 

the embryonal development are prominent, we arrive at the second 
reports of species from America, namely, those contained in 
Linton's (1889) first paper. This was followed by a second (1890), 
containing extensions of the first, and much later by others 
(1891, 1897, 1901 and 1901a) dealing with a variety of forms from 
marine and fresh- water fishes. Further anatomical studies by 
Loennberg (1891), Kraemer (1892), Mattz (1893) and Zernecke (1895) 
lead U3 on to Monticelli's (1892) classification, which was the 
most important since the time of Diesing, altho Perrier (1878) had 
in the meantime published his ideas along that line. The next in 
order is Ariola(s (1896) division of the family "Bothriocephalidae " , 
in which incidentally were yet to be found errors regarding the 
position of the bothria. Beginning with 1894 and continuing to 
1900 there was in progress the publication of Braun's "Cestodes" 
in Brann's "Thierreich" , which is by far the most important work on 
the group since it brings together in a comprehensive manner the 
substance of the most important of the earlier works on the morpho- 
logy as well as the system of the group. One of the first papers 
by the late Dr. Luehe, who was the leading authority on the group 
was that (1896) in which he dealt with the nervious system of 
Ligula. Further study led him to publish a few years later (1899) 
his first classification, which was adopted by Braun (1894-1900). 
In the meantime Loennberg (1897) made important contributions to 
the knowledge of the phylogeny of the parasitic flatworms; while 
Gamble (1896) in the "Cambridge Natural History" and Perrier (1897) 
had erected systems of classification which, however, do not have 
nearly as much in their favor for general acceptance as does that 



7 

by Luehe . In 1900 Ariola brought out his revision of the family 
of the Bothriocephalidae , which, however, waa shown by Luehe 
(Ariola, 1901) to be rather of the nature of a compilation, in- 
volving at the same time several omissions, than a distinct advance 
in our knowledge. Then in 1901 there appeared in Lankester's 
"Treatise on Zoology" Benham's classification of Cestodes which 
professedly follows the earlier works of Railiiet and Blanchard. 
Luehe* s (1902a) revision of the bothriocephalid system comes next 
in order. It is this newer system, only slightly modified in 
1910, that is accepted by the writer with several necessary modifi- 
cations which are dealt with below. 

From 1902 until "Die Susswasser fauna Deutschlands" was 
published the literature on the group consists mostly of papers on 
individual species or mere listings. Spengel's (1905) paper on 
"Die Monozootie der Cestoden" ought, however, to be mentioned, 
since it is one of the latest discussions of a question which occu- 
pied a good deal of the attention of many of the older writers. 
Finally Ward (1910) and the writer (1914 a and b) made the latest 
additions to the American literature, while Stiles and Has sail 
(1912) won the gratitude of the younger workers at least by their 
publication of the section of the "Index- Catalogue of Medical and 
Veterinary Zoology" on "Subjects: Cestoda and Cestodaria" which 
the writer has found of inestimable value in the pursuit of his 
studies . 



3 

EXPLANATION OF TERMS 

Owing to not a little confusion which exists in the 
earliest literature regarding the terms of orientation U3ed for 
the cestode body, the writer wishes to here explain those that 
will be employed in the specific descriptions below. Even much 
later than the time of Diesing (1850) the word "lateral" referred 
to the flat surface of the strobila, while "marginal" was and is 
even yet perfectly clear; but from the standpoint of bilater al 
symmetry both words may mean the same thing. Here they are con- 
sidered to be synonymous and are used to refer to any part that 
is situated in, at or towards the edges of the strobila. On the 
other hand, the work "surficial" is introduced from geology to 
take the place of the German word "f lachen3tandig" which is used 
freely in Luehe's papers to mean that the structures in question 
are situated on the broad, flat surfaces of the chain, that is, 
its dorsoventral faces. The end bearing the scalex is called the 
anterior end and the opposite the posterior, as is customary, 
despite controversies regarding which is which. For the sake of 
brevity the words, "length", "depth" and "breadth" (or "width") 
are used instead of the longer terms, diameters in the longitu- 
dinal, in the dorsoventral and in the transverse directions, re- 
spectively, excepting where the organ in question, e.g., the trans- 
versely elongated cirrus-sac of the Triaenophorinae is so shaped 
that it would be confusing to speak of its obvious length as its 
width. Otherwise the usual terms of orientiation are employed. 



3 

Order PSEUDOPHYLLIDEA Carua 1863, nec Luehe 1910, e .p . 

Polyzootic, Ceatodea with moatly unarmed, 3eldom armed, 
scale* without rostellum, or proboacia formation, excepting in the 
Kaplobothriinae where the primary 3calex ia provided with four 
protusible proboscides resembling those of the Trypanorhyncha; 
usually with two weakly developed sucking grooves, which in indi- 
vidual cases are modified by the strong development of their walls 
or by more or less extensive fusion of their edges, so that they 
may appear funnel-shaped or tubular, which may also unite with 
each other more or less completely to form an unpaired terminal 
adhesive organ, become rudimentary or entirely absent, in which 
latter case they are likewise replaced by a termainal functional 
organ of attachment. Furthermore the development of a paeudo- 
3calex takes place occasionally. Head stalk absent. External 
segmentation more or less pronounced, only seldom completely ab- 
sent. Genitalia in each segment usually single, seldom double. 
Their development proceeds from ahead backwards and does not con- 
tinue to a degeneration of the reproductive glands, but the major- 
ity of the proglottides, being at the same stage of development, 
bring their sexual products to maturity at the same time, so that 
in all of them new eggs are formed continuously and all the eggs 
of the whole animal are at the same stage of embryonic development. 
A surficial opening of the uterus is always present. 

Testes numerous, vas deferens strongly coiled without a 
true seminal vesicle. Ovary near the posterior end of the pro- 
glottis, mostly median in the case of single genitalia, seldom 
approaching the margin of the strobila bearing the genital opening 



(that of the cirrus and vagina). Vitelline follicles very 
numerous, mostly in the cortical, seldom in the medullary paren- 
chyna. Uterus, a more or less winding canal, the individual coils 
of which converge somewhat towards the centre of the proglottis 
to form the so-called rosette; but in other forms it enlarges to 
a capacious cavity, the uterus sac, from which the duct-like be- 
ginning of the uterus is sharply separated. Eggs operculate or 
non-operculate , developing mostly only after being laid, but in 
other cases within the uterus. 

The above diagnosis of the order is that of Luehe (1910: 
11-13) minus the family Caryophyllaeidae and partly emended to 
accommodate the subfamily Haploboth^inae in which what the writer 
considers (vide infra) to be the true (or primary) scolex is de- 
prived of bothria but provided with four eversibie proboscides quite 
comparable in structure to those of the order Trypanorhyncha . As 
to the phylogenetic significance of these organs which are quite 
unique for bothriocephalids, the writer is not yet in a position 
to make any definite statements, since the earliest stages in their 
development have not been observed. It is evident that we can not 
now consider what was formerly (Cooper, 1914 a and 1914 b) called 
the scalex of Haplobothri um to be a true scalex but only the fore- 
most segment of the alult or secondary strobila, which was indeed 
foreshadowed by the writer by his emphasis of its resemblance in- 
ternally as well as externally to the segments immediately following 
Whether or not a pair of bothria were originally present or are 
pres-nt in the very earliest stages, whether such bothria have be- 
come modified into the proboscides, or whether the latter have 



11 

developed from four separate "accessory suckers" (vide infra, 
p. ), as believed by Pintner (1880) to be the case in the 
Trypaaorhyncha , must remain mere suggestions for the present. Fur- 
thermore, as to the formation of segments we have in Kaplobot h rium 
not only conditions quite similar to those in B othriocephalus 
s.str. and other genera in which there is no neck but segmentation 
begins immediately behind the scale*, but those reminding us of the 
proliferation of scalices in echinococcus . In the former case we 
shall see below where the process is described more in detail 
(p. ), that a primary segment divides up into secondary segments, 
these into tertiary segments, and soon until there may be eventually 
32 or more genital segments corresponding to one primary segment 
formed immediately behind the scalex. In Haplobo thrium a primary 
strobila divides up into primary segments, these subdivide into 
secondary segments, the definitive joints of the ordinary strobila 
met with, which in turn may subdivided again and evidently inde- 
finitely to form new chains. The chief difference between these 
two cases is one of degree of regularity in the 3ubdividion. 
Whereas in Bothrio ce phalu s the whole anterior region of the worm is 
affected, evidently no division taking place after the rudiments 
of the reproductive organs have become separated from the common 
rudiment, and the subsegments remain attached to one another, in 
Haploboth rium not only do the primary segments separate as secon- 
dary strobila8, but in the latter only a limited region is in- 
volved in further subdivision. On the other hand there is some- 
what of a resemblance between this manner of subdivision in 
Ha plobot hrium and + hat of the larval Echino coccifer in that the 



12 

atrobilas are developed from an original "nurse". That is, we 
might look upon the primary strobila of the former as a nurse from 
which are developed segments, comparable to the daughter-cysts of 
an echinococcus , which in turn produce (secondary) 3colices and 
eventually strobilaa. In other words we might recognize at first 
sight a sort of alternation of generations in the case of Haplo- 
bothrium. But this comparison is only a superficial one, for 
as will be shown below (under Haplobothriinae) the secondary 
scale* cannot be considered to be a true scalex nor the secondary 
strobila a true strobila; but the primary strobila with its four 
proboscides must be regarded as such. Finally, this peculiar 
method of segmentation reminds one of the asexual budding of some 
of the oligochaete worms, particularly as regards the prolifera- 
tion of subsegments in the anterior region of the first formed 
divisions; but further than this the comparison can scarcely be 
carried . 

Family I. DIPHYLLOBOTHFIIDAE, Luehe 1910, char, emend. 

Polyzootic Pseudophyllidea with unarmed or (seldom) armed 
scalex. Surficial bothia variously developed; they may be modi- 
fied to form sucking tubes, each with an anterior and a posterior 
opening, thru the growth together of their free edges, while an 
unpaired terminal organ of attachment can serve as a funcational 
substitute for the rudimentary bothia or result from the more or 
less complete fusion of both bothia. The whole scalex may be re- 
placed in sexually mature specimens by a pseudoscolex; or it may 
be (Haplobothriinae) provided with four protrusible proboscides. 



13 

Neck present or absent. External segmentation most, present, seldom 
absent. Genital organs numerous, mostly single in each proglottis, 
seldom double. Cirrus unarmed, excepting in Haplocothrium, with 
cleft cuticula. Opening of cirrus and vagina surficial or 
marginal, in the first case always on the same surface as the 
uterus opening and ahead of this as well as always in the median 
line. of the genital complex, also in the median line of the pro- 
glottis in the case of single genitalia. Both surfaces of the 
chain of proglottides, apart from the genital openings, similarly 
3haped. Receptaculum seminis formed by a local enlargement of 
the vagina near its inner end, which as a rule is sharply separated 
from the spermi-duct (terminal portion of the vagina). Uterus, 
a long more or less winding canal, usually in the form of a 
rosette, arising from almost transversely directed coils crossing 
the median line. It may be locally more or les3 enlarged, but 
seldom forms an undivided uterus-sac distinct from the uterine 
duct, as in the Ptychobothriidae . Eggs thick shelled, with oper- 
cula, excepting in the Marsipometrinae; their formation is carried 
on continuously in fully-developed proglottides; embryonal develop- 
ment usually after liberation, seldom in the uterus, in which, 
however, one finds all stages side by side. 
Parasites of vertebrates. 

Luehe's (1910: 16-17) diagnosis is here emended to in- 
clude the new subfamilies Haplobothriinae and Marsipometrinae. 
In the former not only is the scalex radically different from that 
of any other member of the family, but the cirrus is armed with 
minute spines and there is a distinct uterus-sac separate from 



the uterine duct as in the Ptychobothriidae; while in the latter 
there is likewise a uterus-3ac and the eggs are not provided with 
opercula. The scalex of Haplo bothrium is discussed above under 
the order and more at length below under the species so that no 
further mention of it is necessary at this point. The cirrus, 
however, would seem to exclude the genus from the family Ptycho- 
bothriidae as well as from the Diphyllobothriidae , since it is 
not "unbestachelt , mit zerklufteter cuticula", but provided with 
minute yet distinct cuticular spines bearing some resemblance to 
those of the Acanthophallidae (= Amphi tretidae ) as pointed out 
elsewhere by the writer (1914 a : 3) . But H. globuli forme is 
otherwise so nearly related to D. latum that it does not seem 
wise to remove it from the family on this account, especially 
since these 3pines are so minute and since the evidence points 
to their being probably of little, if any, functional importance. 
As regards the uterus, on the other hand, we have something which 
is quite different from that of any of the members of this family 
in that it is distinctly divided into uterine duct and uterus-sac 
as in the Ptychobothriidae . It is true that in the genus 
Scyp hocepha lus one or two of the coils of the uterine rosette be- 
comes much enlarged when the organ is filled with eggs, while in 
Bothridium , as stated by Luehe (1899 : 49),"Der Uterus bildet 
keine Rosettenf orm, la3st jedoch Uteringang and Uterus s.str. 
deutlich unterscheiden; letzterer stellt gewissenmassen eine 
zweitheilige Uterushbhle dar, indem zwei hinter einander gelegene 
gro3se Hohlr'aume durch einen kur^en und diinnen Canal miteinander 
in Verbindung stehen"; but in neither case is there a single 



15 

uterus-sac, distinct and separate from the uterine duct or 
beginning of the uterus but only a modified rosette formation. 
Rohoz (1383 : 282) in describing the development of the uterus 
of Bothridium pithon i s said that: "In dieser Weise ist er natur- 
lich nur in jiingeren Gliedern entwickelt, wahrend er dort, wo die 
Befruchtung schon beendet ist, in Folee der immer starkeren An- 
sammlung von den mit chitinoser Hiille umgebener Eiern immer 
grosser wird und sich schliesslioh zu einem die gange Mi ttelschicht 
ausfullenden Sack ausbreitet . " It would thus seem to be 
comparable to that of the Ptychobothriidae in that its functional 
sac is developed by a distal enlargement of the original duct, 
which gradually encroaches upon the medulla (vide infra), but 
evidently there is no separation of the organ into two distinct 
parts at any stage as there is in Hapl obothr ium . And, as 
emphasized elsewhere by the writer (1914a : 2-3), this separation 
is present at all stages in the development of the organ, which 
as a matter of fact proceeds in quite the same manner as that of 
Bothr i o cephalu s . In Marsipomet ra, on the other hand, even tho 
the sac is formed in the same way, it is never very sharply 
separated from the uterine duct, altho such appears to be the case 
in the adult. Reference to the specific description below will 
elucidate this latter point which seems somewhat paradoxical. 
Finally, as regards the fact that its eggs are not provided with 
opercula, Mars ipometra stands alone. This character would place 
it at once in the Ptychobothriidae, but it is otherwise so closely 
related to the subfamily Triaenophorinae , that the writer here 
emends the lamily to accommodate it. Thus we see that these two 



IS 

isolated genera have such a disturbing influence on our conceptions 
of the two families that the latter now seem to be much more closely 
related than was formerly thought to be the case. 

Subfamily 1. Ligulinae, Luehe 1899 

Scolex unarmed, very 3hort, almost triangular, with 
anterior end, more or less drawn out into a point according to the 
state of contraction, passing directly into the chain of pro- 
glottides or the similarly 3haped unjointed body; surficial bottria 
small, weakly developed. Neck absent. Formation of proglottides 
complete, confined to the anterior end or (in young animals) 
absent. Posterior end rounded. Nerv.' ous system distinguished by 
a large number of plexus forming longitudinal nerves near both 
chief strands. Genital organs in sexually mature individuals com- 
pletely developed close behind the sc&lex. Genital openings sur- 
ficial, ventral, lying behind or near onejanother, near the median 
line. Testes in a simple dorsal layer in the lateral fields of 
the medullary parenchyma, for the most part lateral to the nerve 
strands. Ovary and shell-gland median, the former ventral, the 
latter dorsal. Vitelline follicles in the form of a mantle in 
the lateral fields of the cortical parenchyma. Vas deferens63 
enlarged to a muscular bulb before entering the cirrus- sac. Re- 
ceptaculum seminis large, 3harply separated from the short and 
narrow spermiduct. 

Sexually mature in the intestines of water birds; present 
as larvae in the body-cavitie3 of teleosts vrhere they grow quite 
large and form the rudiments of the reproductive organs; occasion- 



17 

ally al30 observed free in the water which they reach evidently 
by the rupture of the greatly distended bodywall of the interme- 
diate host. 

Type genus: Ligu la Blach. 



In the above diagnosis of the subfamily by Lushe (1910: 
17-18) the statement that the testes are "in einfacher dorsaler 
Schicht den Seitenf eldern des markparenchyms grossentheils 
lateralwarts von den Mark3trangen" is somewhat confusing, for it 
is strictly correct only when the whole number of testes is taken 
into consideration. In transections of both Ligu la and Sohisto- 
ceph alus the nerve strand was actually found by the writer to be 
more than halfway from the median line to the margin of the medulla, 
but the testes were much more closely crowded in the lateral por- 
tion of the field, hence making their total number there more than 
in the median field. But the differences between the two fielda 
on each side in this regard were seen in confirmatory frontal 
sections to be much greater in Lip^ila than in Schistocephalu3 . 

Genus 1 Ligula Bloch, 1783. 

Taenia (part.) Auctorum 

Fasciola (part.) Linnaeus, 1758. 

Faseiola_ (part . ) Linnaeus, 1737. 

Lip;ula Bloch, 1782. 

Fasciola (part.) Goeze, 1782. 
Bothriocephalus (part.) ilitzsch, 1824, 

Ligula Creplin, 1839. 

Difrothrium (part.) Donnadieu, 1877. 



13 

Generic diagnosis: Bothria as well as external segmentation 
completely absent from the larvae, Ooth develop simultaneously with 
the maturation of the sex-organs in the definitive host, and then 
the external segmentation, which does not correspond with the 
internal, is confined to the anterior end. Longitudinal and 
transverse muscles irregularly interwoven in the anterior end, 
posteriorly separated into an inner transverse and an outer longi- 
tudinal layer. 

Type (and only) species: Ligula Intestinal! a (L) . 



Ligula intestinalis (Linnaeus , 1758) 
(Figs. I - e.) 

Larval stage; 



1713 


Taenia 




Geoifroy 


1713 


; 50. 


1740 


Taenia 


capitata 


Frisch 


1740 


; 121. 


1758 


Fasciola intestinalis 


Linnaeus 


1758 


: 649. 


1767 


Fasciola intestinalis 


Linnaeus 


1767 


:I078. 


1781 


Taenia 


cingulum 


Pallas 


1781 


: 95. 


1782 


Ligula 


pisciura 


Bloch 


1782 


: 2. 


1782 


Fasciola abdominalis 


Goeze 


1782 


: 187. 


1790 


Ligula 


abdominalis 


Gmelin 


1790 


:3043. 


1790 


Ligula 


a. cobitidis 


Gmelin 


1790 


:3043. 


1790 


Ligula a.Cyprinorum 


Gmelin 


1790 


:3043. 


1790 


Ligula 


a.alburni 


Gmelin 


1790 


:3043. 


1790 


Ligula 


a. bramae 


Gmelin 


1790 


:3043. 


17 90 


Ligula 


a. carassii 


Gmelin 


1790 


:3043. 


1790 


Ligula 


a. gobionis 


Gmelin 


1790 


:3043. 


17 90 


Ligula 


a. leuscisci 


Gmelin 


1790 


;3043. 


1790 


Ligula 


a. trincae 


Gmelin 


1790 


:3043. 


1790 


Ligula 


a. vimbae 


Gmelin 


1790 


:3043. 


1790 


Ligula 


petromyzontis 


Schrank 


1790 


: 119. 


1793 


Ligula 


salvelini 


Schrank 


1793 


: 143. 


1802 


Ligula 


simplicissima 


Rudolphi 


1802 


: 99. 


1803 


Ligula 


alburni 


Zeder 


1803 


: 266. 


1803 


Ligula 


bramae 


Zeder 


1803 


; 263. 


1803 


Ligula 


carassii 


Zeder 


1803 


: 262-3. 


1803 


Ligula 


cobitidis 


Zeder 


1803 


: 266. 


1803 


Ligula 


colymbi 


Zeder 


1803 


: 266 















SO 


1803 


Ligula 


gobionis 


Zeder 


1803 




265. 


1803 


r Ligula 


leucisci 


Zeder 


1803 


• 


265. 


1803 


Ligula 


tincae 


Xeder 


1803 


• 


265. 


1803 


Li*mla 


virabae 


Zeder 


1803 


* 


265. 


1810 


Ligula 


acuminata 


Rudolphi 


1810 




24. 


1810 


Ligula cingulum 


Kudolpni 


1810 




20-22,31. 


1810 


Ligula 


c ons tringens 


Rudolphi 


1810 


• 


22-24. 


1810 


Ligula 


c ont ortrix 


Rudolphi 


1810 


: 


18-10. 


1819 


Ligula 


simplicissima 


Rudolphi 


1819 




134. 


? 1819 


Ligula 


c r is pa 


Rudolphi 


18189: 


134-35. 


1818 


Ligula 


edulis 


Briganti 


1819 




209. 


1839 


Ligula 


simp lie is s ima 


Creplin 


1839 


! 


295. 


1839 


Ligula 


monograrama 


Creplin 


183 9 


: 


296. 


1839 


Ligula 


digramraa 


Creplin 


1839 


• 


296. 


1853 


Ligula 


simplicissima 


Baird 


1853 


* 


95. 


1855 


Ligula 


monogramma 


Leidy 


1855 




444. 


1861 


Ligula 


monogramma 


Beneden 


1862 




139. 


1891 


Ligula 


catostomi 


Linton 


TOOT 

1891 




66. 


1896 


Ligula 


monogramma 


Zschokke 


1896 




773,774,775. 


1897 


Bibothrium ligula 


Linton 


1897 




438. 


1899 


Ligula 


abdorainalis 


Luehe 


1899 




52. 


Adult 


stage: 












1782 


Ligula 


avium 


Bloch 


1782 




4. 


1783 


Pasciola intestinalis 


Goeze 


1782 




183. 


1790 


Ligula 


intestinalis 


Graelin 


1790 




3042. 


1802 


Ligula 


simplicissima 


Kudolphi 


1802 




99. 


1803 


Ligula 


colymbi 


Zeder 


1803 




266 • 


1810 


Ligula 


uniserialis 


Rudolphi 


1810 




12. 



1810 


Ligula 


alternans 


Rudolphi 


1810 


; 


13. 


1810 


Ligula 


interrupta 


Rudolphi 


1310 


: 


15. 


1810 


Ligula 


sparsa 


Rudolphi 


1810 


• 


16. 


1819 


Ligula 


uniserialis 


Rudolphi 


1819 


■ 


132. 


1819 


Ligula 


alternans 


Rudolphi 


1819 




133. 


1819 


Ligula 


interrupta 


Rudolphi 


1819 




133. 


1819 


Ligula 


sparsa 


Rudolphi 


1819 


; 


133. 


1824 


Bothriocephalic semi- 
ligula. 


Nitzsch 


1824 




98 


1839 


Ligula 


uniserialis 


Creplin 


1839 


• 


296. 


1839 


Ligula 


interrupta 


Creplin 


±859 


• 


296. 


1844 


Ligula 


sparsa 


Bellingham 


1844 


• 


165. 


1845 


Ligula 


uniserialis 


Dujardin 


1845 


• 


628. 


1845 


Ligula 


alternans 


Dujardin 


1845 


• 


629. 


1845 


Ligula 


interrupta 


Dujardin 


1845 


• 


629. 


1845 


Ligula 


sparsa 


Dujardin 


1845 


: 


629. 


1845 


Ligula 


nodosa 


Dujardin 


1845 


: 


629. 


1850 


Ligula 


monogramraa 


Diesing 


1850 


• 


579. 


1850 


Ligula 


digramma 


Diesing 


1850 


■ 


580. 


1853 


Ligula 


interrupta 


Baird 


1853 


: 


96. 


1853 


Ligula 


sparsa 


Baird 


1853 


: 


96. 


1854 


Ligula 


monogramraa 


Diesing 


1854 


: 


18. 


1854 


Ligula 


digramma 


Diesing 


1854 


• 


18. 


1856 


Ligula 


reptans 


Leidy 


1856 


• 


46. 


1863 


Ligula 


monogramraa 


Diesing 


1865 




2o0. 


1863 


Ligula digramma 


Diesing 


1863 




231. 


1870 


Ligula 


monogramma 


Willeraoes-Suhm 1870; 


94. 


1877 


Dibothrium ligula 


Donnadieu 


1877 




495. 


1881 


Ligula 


simplicissima 


Moniez 


1881 




37,8 



22 



1882 


ligula 


simplicissima 


Kiessiing 


1882. 




1884 


Dioothrium ligula 


Zschokke 


1884 : 


26. 


1885 


Ligula 


sirapiicissima 


Schauinsland 


1885 ; 


550. 


1888 


Ligula 


simpliciss iraa 


Memiec 


1888 : 


2. 


1893 


Ligula 


monogramma 


Blsson ; 


1893 : 


15. 


1894 


Ligula 


simplicissima 


Stiles &Hass 


all 1894 


:331. 


1895 


Ligula monogramma 


Zernecke 


1895 ; 


93. 


1895 


Ligula 


digramma 


Zernecke 


1895 ; 


93. 


1896 


Ligula 


simplicissima 


Zschokke 


1896 ; 


773,774,775. 


1898 


Ligula digramma 


Cohn 


£898 : 


134. 


1898 


Ligula 


uniserialis 


Luehe 


1898 ; 


286. 


1898 


Ligula 


uniserialis 


Muehling 


18 98 : 


32. 


1898 


Ligula 


monogramma 


Stossich 


1898 : 


118. 


1899 


Ligula 


intestinalis 


Luehe 


1899 : 


52. 


1900 


Ligula 


avium 


Braun 


1900 : 


1687. 


3.900 


Ligula 


uniserialis 


Wolffhuegel 


1900 : 


63. 


1901 


Ligula 


intestinalis 


Linstow 


1901. 




1902 


Ligula 


monogramma 


Parona 


1902 : 


7. 


1902 


Ligula 


intestinalis 


Schneider 


1903a: 


13. 




Ligula 


intestinalis 


Lins t ow 


I90o : 


<s0. 


1910 


Ligula 


intestinalis 


Luehe 


1910 : 


18. 



Specific diagnosis; with the characters of the genus. Large 
worms from 100 to lOOOmra.in length by 5 to 15 in breadth. Anter- 
ior end rounded, protruding, bothria faint. Strobila greatly elon- 
gate, depressed, maximum breadth anterior to the middle , gradually 
tapering to the posterior end. Body crossed by irregular ridges 
and furrows, and wavy at the margins in the adult with 35 to 40 
external segments anteriorly. Deep median groove on each surface 



S3 

in the larva, two very shallow parallel grooves near the median 
line on the dorsal surface in the adult. 

Cuticuia 5 - 20p. in thickness , sub cuticuia 70 -100. ixerve 
strands 50^ in diameter. Excretory vessels numerous in two lay- 
ers, one close beneath the vitelline glands ( cortical) another a- 
mong the main body muscles. 

Genitalia from 0.10 to 0. 20mm. apart . Genital cloaca a nar- 
rov transverse slit ,0.18-0.20 x 0.02 - 0.05mm. into which open 
separately the cirrus , uterus and vagina, the latter constantly be- 
tween the other two which alternate irregularly from side to, side. 

Testes in single dorsal layer in the medulla, interrupted only 
medially, 35 to 40 in transection, 115 x 85 x 45^ in dimensions. 
Vas deferens up to 30 (a. in diameter , loosely coiled above the cirrus 
sac. Seminal vesicle small, close above the latter, 65 - 80 x 40 - 
65|^ . Cirrus-sac somewhat lateral, ovoid, with thin walls , 185-200 
x 130-145 x 130-135 u~ . Cirrus proper within cirrus-sac , long and 
coiled, 2by, in diameter. 

Vagina 15 to 30(ll in diameter , receptaculum serainis 75 p. . 
Sperraiduct short, 25 x 12 ^ . Ovary 1.5mm. in diameter; wing great 
ly depressed, isthmus prominent and not in the median liAe but al- 
ternating irregularly from side to side opposite the cirrus-sac; 
ova in same 15 x 13 fv . Oocapt 18-80 ^ in diameter, oviduct 15-20 ju/. 
Vitelline reservoir ellipsoidal in shape, 50-60 x 30^, in a layer 
close beneath the subcuticuia and broken only ventrally. Shell- 
gland composed of much elongated cells with enlarged bodies and 
narrow necks connecting with the oviduct for 30^Lof its length. 
Uterus a mass of coils in the median line, 0. 5-0, 6mm. im diameter, 
that of the tube being 60 ^. 













34 


Eg 


gs, 49-65 x 


31-42^ . 








Habitat : As 


larvae in the body-cavities 


of teleosti 


s; adults 


in the 


intestines 


of wading and diving birds. 






Host 

Leuciscus vulgaris 


Locality 


Collector 


Authority 




Pa 1 1 a c 
id Xiao 


Diesing 


1850:581. 




rutilis 




Hubner 




• 


N 


ii 




Z- O V^ilUiVi\C 


Zschokke 


1884:26. 


H 


ii 


R ncci f ton 


Ti/ii i o Vi 1 "i n rr 
MU t: X 1 X X lig 


Muehling 


1898:33. 


H 


ii 


Orlja-See,Pe- 


Birulja 


Linstow 


1903:285. 


!1 


ii 


i-Strabad Bay, 
Caspian Saa. 


iVia x imovi c 

AflWt ^VJb. IUU V X W' 


ii 


ii ii 


H 


leuciscus 






Luehe 


1910:19. 


H 


ervtiiro- 

ohthalraus 




Brems er 


Diesing 


1850:581. 


if 


puicneij.u5 




Baird 


Leidy 


1855:444. 


II 


phoxinus 


L.Stors jon, 
Jeratland,and 
Bonan, Sweden. 


Qlsson 


Olsson 


1893:15. 


Abrami 


s blicca 




Goeze 


Diesing 


1850:581. 


ii 


brania 




ivuu. uxpzii. 


ii 


ii ii 


it 


ii 




R v» o m o d 

Dreniser 


N 


ii it 


ii 


ii 




lvl,Dl6DO XQ. , 

Coll. Brit. 
Museum. 


Baird 


1853:95. 


it 


ii 


L.Finjasjon, 
Scania,Sv. eden 


Olsson 


Olsson 


1893:15. 


ii 


ii 




Zschokke 


Zschokke 


1896:775. 


•i 


ii 


East Prussia 


Schauinsland 


wuehling 


1898:33. 


ii 


ii 


Ladoga-See 


Firlei 


Liastov. 


1903:285. 


it 


ii 


Astrachan 


Baer 


ii 


ii ii 



■ bramis vimba 


Petrograd 


Prichodko 


Linstow 


.15 

1903 :285. 


" b,i orkna 






Luehe 


1910: 19. 






Cyprinus Jar&j&a 




Bloch 


Bloch 


1782 : 2 . 






Borke 


Goeze 


1783:187. 


■CyRHiaus ^obio 




Diocn 


Bloch 




Cvprinus alburnus 




it 


ii 


it ii 


It II 


Berlin 


Goeze 


Goeze 


1782 : 187. 


Cyprinus tines 




Bonnet 


Bloch 


1782 :2 . 


" blicca 






Rudolphi 


1810:19. 






" carassius 




Pallas 




H 22 . 






Creplin 


Creplin 


1839:^96. 


" lacustris 


Italy 


origanti 


Donnadieu 


1877 :339. 


" leuciscus 


La Reole & 

Ki oil f-PnYi e 

is eui i ons 


Q,ueyron 
Bremser 


Queyron 
Diesing 
Linstov. 


1905:CVI. 
1850:581,4 
1903:285. 


" rapax 


Iljmen-See, 
Kovogorod 


Varpachov- 
ski j 


Qofrio vulgaris 




Rudolphi 


Diesing 


1850: 581, 


" fluviatilis 


Basel 


Zschokke 


Zschokke 


1896 :774. 




Rossitten 


inuehling 


Miehiinfe 


1898:33. 


it it 


Angara -Fluss, 
Siberia 


Cekanovski j 


Linstow 


1905:285. 


Carassius gibelio 




Rudolphi 


Diesing 


1850:581. 


" vjilgaxis 




Diesing 




w :579. 




Saratov 




Linstow 


1903 :285. 


" carassius 






Luehe 


1910:19. 


Amoetes branchialis 




Schrank 


Diesing 


1850:581. 


Petromyzon branchia 
lis 


i- 


Schrank 


Rudolphi 


1810:24. 


Cobitis taenia 




Frisch 




1819:134. 


ii ii 




Bloch 


ii 


it it 











<~>o 


Coxitis aculeata 




Bloch 


Bloch 


1782 ;2 . 


Rnlmn 5=?fa 1 vel inup 




Schrank 


Diesing 


1850: 581. 








• 


£j "juris glanjs 




Mus. Vienna 




• 


Esox lucius 










ii n 




Olsson 


Olsson 


1895:15. 


Perca fluviatilis 




mus .Vienna 


Diesing 


1850:581. 


ii it 


L.St or sj on, 
J smt land , and 
Bbnan,Sv. eden 


Olsson 


Olsson 


1893:15. 


ULCi.operca s_suodxa 




Mus .Vienna 


Diesing 


1850:581. 


" lucioperca 






Luehe 


1910:19. 






? Phoca vitulina 


Berlin 


Rudolphi 


Rudolphi 


1819:135. 


Morrhua americana 




Schaf irt 


Leidy 


1855:44-4. 


Squalius cephalus 




Zschokke 


Zschokke 


1884:26. 


" turcicus 


l scnaidyr-goi 
See , Armenia 


Brandt 


Linstow 


1903:285. 


Alburnus lucidus 




Zschokke 


Zschokke 


1884:26. 


it ii 


Jedwabno 


Braun 


Muehling 


1898:33. 


u ii 


Langviken Bay, 
Finland • 


Levander 


Schneider 


1902a: 13. 


" alburnus 






Luehe 


1910:19. 






Atherina raocho 


Cagliari 


Parona and 
ivuazza 


Par .(iMazz 


.1900:233. 


B lice a bj orkna 




Linstow 


Linstow 


1901:629. 


Catostomus ardens 


Yell. Nat. Park 


D .S.Jordan 


Linton 


1891:65. 


Chondrostoraa nas- 
us 


Basel 


Zschokke 


Zschokke 


1896:775. 


Catostomus lati- 
pinnis 


;Jila R. and 
Salt R., Ari- 
zona 


E. Palmer 


Linton 


1897:438. 


Osmerus mordax Potomac R. . 

H&gerstown,ivid. 


C .L.Riden- 
our 


it 


it ii 



37 

Hybognathus nucha lis Linton 1897:458. 

^o^ropis cornutus Fourth L., F.Mather M " N 

Adirondacks , 
Nev York 

Sch^zopv^Qpsis Tan-la-Gebirge, Przevalskij Linstow 1903:285 

KQZ.kQyJi Tibet 

ftjemachilus strau - Issyk-kul-See , P. Schmidt " " " 

chi Bai Karasu 

" » Przevalskij- " " " " 

Bai, Aral Sea , „ 

Tinea vulgaris - • Neveu- 

Lemaire lfioa :88. 

Gobio gobjo Luehe 1910 : 19. 

Scardinius ery- » " " 

thr ophthalmus 

Ameiurus sp. Charlevoix, H?B.Ward Cooper 

Michigan (the present paper) 

Alosa ohiensis Keokuk, Iowa " " 

Perca fluvia tiiis Walnut L. ,Mich. M " 

" f lavescens Go-Home Bay,Mus- Cooper " 
koka District, 
Ontario. 

Catostomus commer- Walnut L. ,Mich. H.B.Ward " 
§ Qai i 

" ■ Turtle L.,Mich. H.R.Hill » 

JLotropis cornutus Go-Home Bay Cooper " 

" cavuga Turtle Lake H.R.Hill » 

Micropterus djlip- Go-Home Bay Cooper " 

miau 

Amblot)lites rupes- " " " '* 

13? — 

Gasterosteus bispi - Chamcook L,N ew " " 
nosus " atTInsii Brunswick 

Free on Shore Turtle L. .Mich. H.h.uill " 



28 




braun 
Bremser 
Mus .Vienna 
Creplin 
Braun 

Hildebrandt 



Diesing 

Rudolphi 

Creplin 

Rudolphi 

Diesing 

Luehe 



Hildebrandt Rudolphi 
Mus. Vienna " 
M.C.V. Diesing 



blovcov 

M.C.V. 

mus. Vienna 
ii ii 

Mehlis 
Hubner 
Bloch 
Rudolphi 

Mus. Vienna 

NitEseh 

Mus. Vienna 
Mehlis 



Linstov- 

Diesing 

Rudolphi 
ti 

Diesing 

Kudolphi 

Bloch 

Stiles & 
Hassan 

Rudolphi 

Luehe 

Mtzsch 

Rudolphi 

Diesing 

Luehe 



1850:580. 
ii ii 

1819:133. 

1839:296, 

1810:12. 

1850:589. 

1910:18. 
a ii 

1810:16. 

1819:133, 

1850:580, 
ii ii 

1903:284, 

1850:580, 
1819 :13o, 
" :134, 
1850 :581< 

1810:15. 

1782:4. 

1912:266, 
1819;134, 
1910:18. 
1824:98. 
1819 :134, 
1850:581, 
109ft: 286 













90 


P.ihlvmbns suhcrista 




uus. Vienna 


Rudolphi 
Diesing 


1819:134. 


Pod ic 


epjk au.r\tu,s, 




Bloch 


leou : ooi. 


M 


II 




Hubner 






tl 


II 


Re nnes, France 


Dujardin 


Dujardin 


1845 :629. 


It 


•1 




M.C .V, 


Diesing 


1850: 580. 


It 


II 




ivi.Siebold, Baird 
Coll. Br. Mus. 


1853:96. 


II 


II 


Ireland 


Bellingham 


Belling- 
ham 


1844:165. 


II 


II 


Koenigsberg 


Braun 


wiuehling 


1898:32. 


II 


II 


Pillau 


wiuehling 


ii 


it it 

• 


|| 


II 


Trieste 


Stossich 


Stossich 


1898:118. 


|| 


It 




Wolffhugel 


Wolffhugell900:62. 






• ~ ~ ~ * ■ 


Lins t ov. 


Lins t ow 


1901:629. 




II 


Venice 


Stossich 


Parona 


1902;7. 


II 


II 


Varese 


Parona 


ii 


n ii 


II 




ivionfalcone 


St ossich 


Stossich 


1898:118. 


II 


nitric olli 


s Trieste 




ii 


ii ii 


II 


rubricollis 


M.C.V. 


Diesing 


1850:580. 


II 




Trieste 


Stossich 


Stossich 


1898:118. 


An, as 


boschas fera 




Mus .Vienna 


Rudolphi 


1819:134. 


ii 


boschas 






Luehe 


1910:18. 










ar-entatus' 


Ciryphswald 


Schilling 


Diesing 


1854:19. 


11 


ii 


Kainsk,Enisse j 


Middendorf f 


Lins tow 


1905:20, 


H 


©anus 




Bremser 


Rudolphi 


1810:13. 


II 


melanoceph- 


Varese 


Parona 


Parona 


1902:7. 




(alus 










II 


parasiticus 




Bremser 


Rudolphi 


1810:13. 


II 


•i 




Mus .Vienna 


it 


1819:135. 











30 


Larus pelecanus 
carbonis 




Mus .Vienna 


Rudolphi 


1819:133. 


Larus D.vgmaei 








• 


" ridibundus 




Bremser 




1810:13. 




Rossitten 


wiuehling 


Muehiing 


1898:33. 


- 

n tridactylus 




Hufaner 


Rudolphi 


1810:13. 


sterna .hi r undo 




Mus .Vienna 




1819:133. 


" nigra 








• • 


Fergus aJLbel^us 




Bloch 


Bloch 


1782 :4. 






M.C .V. 


Diesing 


1850:581. 


" merganser 




Kitzsch 


Rudolphi 


1819:133. 






Bloch 


Bloch 


1782:4. 




Belgium 


Beneden 


Diesing 


1863:231. 






Linstow 


Linstor 


1901:629. 


" minutus 






Gmelin 17 90:3042. 






■ serrator 




M.C .V. 


Diesing 


1850:581. 






wuehiing 


wiuehling 


1898:33; 




Gulf of Finland 


Schneider 


Schneider 


1902a: 13. 


„ 

fl^cticorax sp. 




Mus. Vienna 


Rudolphi 


1819:133. 








Luehe 
Baird 


1910:18. 
1853:96 


Graculus carbo 




M.Siebold, 
Coll. Br. Mus 


Fuligula clangula 


Jemtland 


Olsson 


Olsson 


1893:15. 


,Xema minutum 


Trieste 


Stossich 


Stossich 


18 98:118. 


" ridibundum 




Wolf fringe 1 


Wolf f huge 


1 1900:63. 


Urinator arcticus 






Luehe 


1900:18. 






H stellatus 






it 


ii ti 










Rissa tridacty.la 

Stercorarius para- 
sitica 






ii 


ii ii 






N 


it ii 











31 



Hvdrochelidon — Luene 1910: 28 

nigra 



H erodias alba 
rialiaet.us ^ikiciUa 
Aguj.i ,a chrvsaetus 
Corvus .c.Qrpix 



Podilymbus H.B.Ward Cooper 

podice r>s (the present paper) 

Merganser sp % Urbana, Illinois " " 

Col yqbus ho elboelli " " 



32 

As indicated in the aoove synonymipthe greatest confusion 
existed in connection with this species even from the time of 
Linnaeus, all of the older writers recognizing at least two species, 
the larval and the adult, and many, several species under each of 
these. Rudolphi (1810), for instance, accepted four species of 
the former, "ovariis occultatis", and the same number of the latter, 
parasitic in the intestines of birds, "ovariis distinctis". In 
his Entozoorum Synopsis (1819) he reduced the number of larval 
species to two, but retained the 3ame four adult form3 as before. 
The next important move in a systematic direction was by Creplin 
(1839) who divided Rudolphi ' s L. simplicissima into two larval 
species, viz., L. monogramma and L. di gra mma, corresponding re- 
spectively to the previously known L. uniserialis and L. interrupta 
(or alt e mans ) , which plan was followed by Die3ing (1850, 1854, 
and 1863) while Dujardin (1845) and Baird (1853) followed Rudolphi. 
Diesing (1850 : 581) erected a third species, L. rep tan s, to accom- 
modate numerous forms found encysted in the muscles and connective 
tissues of amphibians, reptiles, birds and mammals; but, as pointed 
out by Janicki (1906 : 519-530) several larval species were probably 
included under this heading. Those from avian hosts (i.e.: 583) 
are not given above since they were found only among the muscles and 
under the skin, where L. intestin al! s has never been found in 
birds, so far as the available records indicate. Luehe (1910 : 18) 
did not include them in his list of hosts for thejadult stage of 
the 3pecies. 

Next in order of importance came DonnadieiL* s (1877) 
classical experiments in which,after completely reviewing the 



33 

literature up to date, he conclusively proved that the form found 
in the body cavities of various "bony fishes is the larval stage of 
that present in the intestines of birds. As a result of his 
work he combined the two forms under a new name, Dibot hrium ligula, 
confusing at the same time Schi stocephalus s olid us with Lig?u la 
intes tinali3 . The life-history of the species was later studied 
by Riehm (1833) by feeding methods. Moniez (1331: 37, 31) was the 
first writer to 3tudy the histology of the species, while Kies3ling 
(1833) gave us the first description of it3 general anatomy. As 
emphasised, however, by Linstow (1901), Kiessling^ work is not 
very specific, 3ince he almost constantly disposed of L. intes- 
tinalis by 3aying that in it conditions were the same as in Sen. 
Solidu a. While, apart from Donnadieu and the earlier ?;riters, 
Willemoes-Suhm (1870 : 94) was the first to study the development 
of the embryo with attention to detail. Schauin3land (1835 : 550) 
enlarged upon his observations and gave a more or less complete 
description of the process up to the time of the escape of the 
ciliated larva. Niemiec (1838 : 2 ) described the nerv.ous system, 
and Cohn (1898 : 134) pointed out its resemblances to Sen , solidus 
in this regard. Zernecke (1895) in the meantime dealt in his well 
known work on the finer structure of cestodes with the parenchyma 
and the nerv..ous and muscular 3y3tems in particular, 3ince when 
little has been done in that connection. The question of segmenta- 
tion was studied by Luehe (1898); and later the same writer (1899: 
52) placed the species in his first classification, stating his 
belief that there is only one species of Ligula, viz., L. intes- 
tinal! s (1.). The latter conclusion was also arrived at by 



34 

Linstow (1901 : 638), altho he attributed the specific name to 
Gocze; while in his latest classification Luehe (1910) maintained 
the same view. 

Consequently, taking for granted in the absence of Euro- 
pean material for comparison, that the latter has been established 
as a fact for the European forms, it is for the present writer to 
determine whether we have here in America the same species. And 
so far as the majority of specific characters are concerned, we 
have to rely on the descriptions of Kie3sling and Linstow (1901) 
who seem to have been the only writers to attend to the details of 
the reproductive system, and, a3 mentioned above, Kiesaling's is 
quite inadequate in this connection. The only American reports 
of the species are of larval forms : L. mono gramma by Leidy (1855 : 
444) and Dibothrium ligul a by Linton (1897 : 438), the former 
having also listed (1856 : 46) the doubtful L. repta ns . 

Luehe (1910 : 18) gave the dimensions of the species as 
100-400mm. (occasionally 1 metre) in length by from 5 to 15mm. in 
breadth, not distinguishing, however, between the larva and the 
adult in this regard. Linstow (1901 : 629) reported a larva from 
Blisca bjorkna 200mm. long, 9 broad and 3.5 thick, adult 3 from 
ypdioepe cri status and Mergasser merganser being 150mm. long, 4 
broad and 1.5 thick; and concerning these differences 3aid that: 
"Wenn man die Geschlechtsf orm aus Vo'geln oft kleiner finlet als die 
Larve aus Fischen, so mag das seinen Grund darin haben, dass die 
letztere sich in der GrSsse ihren Wirth anpa3st; die grossen Larven 
in gro33en Fischen k'onnen aber nicht von kleineren Vo'geln ver- 
schlungen werden." The largest larval specimen at hand was one 



35 

from Catostomus oommeraonil whioh measured 425mm. in length by 
15 in maximum breadth, but the largest adult from Merganser Bp. 
wa9 only 142mm. by 7.5. In the larva the anterior end is some- 
what bluntly rounded (Fig. 1) the bothria being visible as very 
short grooves pa33ing over the tip, while in the adult they are 
more elongated and distinct, the end of the strobila being some- 
what protruded, aa shown in Fig. 2. On each surface of the 
larva there is a deep, median, longitudinal furrow, which, however, 
becomes obliterated in the adult, excepting anteriorly, by the 
growth of the reproductive organs, the ducts of which are con- 
fined to the median line of the strobila. When these are developed 
the 3trobila i3 characterized dorsally by a low median ridge 
bounded on each side by a quite shallow groove, and ventrally by 
a greater thickening of the median line, not separated, however, 
by any grooves from the lateral regions. The whole strobila 
gradually tapers from a short distance behind the anterior end, 
where the maximum breadth is located, to the posterior end. 
Whereas in the larva it is quite thick, in the adult it is thin 
asd leaf-like, the margins usually appearing, especially posterior- 
ly, wavy in alcoholic specimens. A pseudosegmentation is present 
in the anterior end of the strobila, but a9 has been known, espe- 
cially since Luehe (1898) emphasized the fact, this division of 
the strobila into segments does not correspond with the internal 
division into true proglottides. Gemmill (1909 : 11) counted 
about 50 of them in the anterior third of the worm, the writer 
38 or 39 for a distance of 13mm . from the tip of one adult speci- 
men (Fig. 2) and 36 for 10mm. in another. They vary considerably 



36 



in length and are often incomplete medially. From the anterior 
region showing external segmentation to the posterior end both 
larvae and adults, but particularly the latter, are crossed by 
very numerous irregular grooves which give the worm its charac- 
teristic appearance, apart from the general shape, as contrasted, 
for instance, with the closely related Sohi stocephalus 3olidus 
(vide infra). The smallest larva met with was one from a small 
specimen of Mioropterus dolomien, 47mm. in length. It gave the 
following measurements: length, 4.9mm.; maximum width, 0.54; width 
one third the length from the anterior end, 0.54, two-thirds, 
0.37; length of bothrial groove, ? 0.07mm. All sizes from this 
to the largest mentioned above were at hand. 

The enticula was "found to have a thickness of from 5 to 
lOp, compared with 16-18 ^ by Kiessling and 2.1^ ( \ ) by Linstow. 
It was seen to be homogeneous in the sections made rather than 
divisible into the three layers described by the former, with 
some tendency, however, for the outer one-quarter to one-sixth to 
take the stain much less than the remainder of the tissue, which 
outer clearer area is often bounded by a very delicate pseudo- 
ciliated layer. There was found to be as much variation not only 
in the thickness of the enticula but also in its structure, that 
nothing more will be said excepting that these remarks apply to 
the larva as well as to the adult. The subcuticula varies from 
70 to SO , — 33-49 pi according to Kiessling and 114 |u to 
Linstow. Calcareous bodies in the characteristically fine paren- 
chyma, described by Moxii ez and Zernecke, and given dimensions of 
13 x 7.8 ju by Linstow, were not observed to the writer's satisfac- 
tion. 



37 

The musculature has been well described histologically 
by Zernecke (1895); while Kiessling 3poke rather briefly of its 
arrangement in the late larva. Later Luehe dealt with the system 
in general (1897a and 1898) and its relation to the nervous 
3y3tem (1896 ), and Lin3tow (1901 ), gave a concise account of 
its arrangement. The writer has nothing of importance to add to 
the contributions of these authors. 

The chief nerve 3trands were found in transections between 
the lateral and median quarters of the transverse diameter of the 
atrobila, in the median frontal plane that is below the neighoor- 
ing testes, and with a diameter of 50 p. . The details of the 
3ystem have been studied by Moniez (1881), Niemiec (1838), 
Zernfccke (1895) and Cohn (1898), the latter of whom found condi- 
tions pretty much as in Schistocephalus , namely ) that each chief 
strand has associated with it six collateral strands, arranged in 
three groups of two each. 

The excretory system was studied by Moniez and Zernecke 
in considerable detail. As pointed out by Lin3tow two regions 
accommodating numerous longitudinal ves3e(3 are present: (1) an 
outer, close beneath the vitelline glands, and (3) an inner, be- 
tween the inner longitudinal and transverse muscles, or as Linstow 
figured, between the former themselves. In the material 3tudied 
the outer-plexus showed very plainly, but the inner vessels were 
found among both sets of muscles and not nearly as distinct as 
the outer one3. 

The sets of genitalia, beginning about IOtmh. from the 
anterior end and very closely crowded together in ths longitudinal 



38 

direction, were found to lie from 0.14 to 0.18mm. apart, 0.13- 
0.15mm. being the data given by Linstow. The openings are 
usually almost exactly in a transverse line, but the cirrus and 
uterus openings alternate irregularly from side to 3ide, that of 
the vagina being constantly in the middle. This alternation of 
the openings is due to the similar altsrnation of the internal 
organs and evidently was the basis upon which the earlier species 
L. di gramma and altern ans were established. The genital cloaca 
is a quite irregular transverse depression, 0.18-0. 30mm, in width 
and ©.03-0. 03mm. in length, the respective measurements by Linstow 
being 0.105 and 0.036mm. 

"The testes lie in a single row, which is only interrupted 
by the uterus, on the dorsal side of the medulla ... n . They are 
from 35 to 40 in number in transections, ellipsoidal in shape, 
their greatest diameters being transverse, a3 indicated by the 
maximum width, length, and depth being, respectively, 115, 45, and 
85 1^ . Linstow gave them as 150 to 180^ long by 88 to 156 ja- 
wide. The loosely coiled vas deferens i3 situated above the cirrus- 
sac (Fig. 3) and is roughly divided into two parts by the laterial 
coil3 of the uterus, one part being immediately above the cirrus- 
sac and the other close to the dorsal body-wall. The duct attains 
a diameter of 30 f*» when filled with spermatozoa. Distally it ex- 
pands into the very small (a3 compared to that of Sch . soli dus) 
seminal vesicle, situated close to the dorsal wall of the cirrus- 
sac. The vesicle was found to be from 65 to 80^ in length by 
40-65 in diameter (156 x 86 ^ , Linstow), oval in shape, the 
narrower end towards the cirrus pouch, and provided with only a 



39 

comparatively fseble musculature. The wall of the structure is 
richly supplied both internally and externally with nuclei which 
are respectively those of the lining epithelium and the myoblast*, 
as in Sc'nist ocephalus (vide infra). The epithelium 13 strongly 
ciliated. The cirrus- sac (Fig. 3) is an ovoid body, somewhat 
flattened dorso-ventrally and obliquely by the uterus and alterna- 
ting irregularly from right to left, always occupying the opposite 
3ide of the median line from the ovarian isthmus and the neighbor- 
ing female ducts. Its wall is quite thin, while apart from the 
cirrus proper which occupied the distal two-thirds, the contents 
consist of loose parenchyma and few retractor mu3cle3. The measure- 
ments of the organ in sections are: dorsoventral diameter, 185-300^.; 
width 130-145; and length, 130-135; which are quite at variance 
with Linstow's diameter of 53 ^ of what he described as a spherical 
organ. Within the cirrus-3ac the vas deferens is not sharply 
separated into ejaculatory duct and cirrus proper, altho the latter 
is quite distinct, closely coiled and as much as 35 in diameter. 

The vagina opens into the common genital cloaca, if one 
may use that name for the depression mentioned above, in the median 
line and usually equidistant from the openings of the cirrus and 
uterus. It paases dorsally thru the cortex and the musculature 
with almost a straight course and then within the medulla turns 
sharply posterolater -ally , in which portion of its course it ha3 a 
diameter of from 15 to 30^ (5pt , Linstow) . Its thin lining of 
cuticula, directly continuous with that of the genital depression, 
gradually passes into a nucleated epithelium, in which no distinct 
cell boundaries were made out, just within the cortex. Dorsal to 



40 



the ovarian isthmus it enlarges into an elongated recei: taoulum 
seminis whichjwas found to have a maximum diameter of 75jul . Lin- 
stow described a spindle-shaped serminal receptacle, 13|<a- in 
diameter, and an oocapt as follows: "dorsal vorider Vereinigung K s tel- 
le der beiden Keimstocksf lugel liegt der ovale, 0.088mm. lange 
und 0.066mm. breite Schluckapparat each of which, however, in 
comparison with that described here by the writer and for Sch . 
sol idus below, seems to be confused with the other. At least the 
oocapt of none of the bothriocephalids studied was found relatively 
so large as indicatec by Linstow in his measurements and in his 
figure, nor was the receptaculum a3 spindle-shaped as shown in the 
latter. In this connection Kie3sling described a swelling of the 
vagina, 46 |a- in diameter, which contained spermatozoa. The spermi- 
duct is so 3hort and of such a small calibre th^t it is quite 
difficult to locate it in sections. It unites with the oviduct a 
short distance from the oocapt (Fig. 4) much as in Sch. solid us, 
after pursuing a horizontal course. It is about 35^ in length 
and 12 ^ in diameter. The ovary is asymmetrical, as stated by 
Kiessling but denied by Linstow, 3ince it consists of a much de- 
pressed lateral wing, situated close to the ventral musculature 
(Fig. 3) and a more median enlarged portion which functions as the 
isthmus in that the oviduct ari3e3 from it. This isthmus-like 
region is not in the median line but about 0.35 K from it, the whole 
organ alternating irregularly from right to left, constantly op- 
posing the cirrus-sac on the other side. It varies from 0.55 to 
0.64mm. in width and has a length laterally of 0.13mm. Its un- 
usual situation is evidently due to the closely crowded condition 



41 

cf the reproductive organs and the pressure exerted by the large 
uterus in the median line. Whereas the wing has a maximum dorso- 
ventral diameter of aoout 60 p- , the isthmus is about 95^ in 
depth and roughly ellipsoidal in 3hape, protruding in sections 
from the dorsal region of the junction of the wing (Fig. 3). Ova 
from the i3thmus were found to "be oval in shape and 15 x 13 jia. in 
size (13-16 , Linstow) . The oocapt is directed horizontally away 
from the side of the isthmus and from the median line. Its dia- 
meter is 18-20^ , with which compare the dimensions of 88 x 56^ 
given by Linstow (vide 3Upra). The oviduct has a diameter of from 
15 to SO p. . Taking a general dorsal course after being joined 
by the spermiduct, it soonEreceives the common vitelline duct 
(Fig. 4) which has only a limited enlargement from a previous dia- 
meter of 10-30 to 30 to form the vitelline reservoir which is 
located close to the oviduct with a length of 40jj~ . The vitelline 
follicles are situated in a layer close beneath the 3ubcutieula 
and are continuous, excepting in the median ventral line. The 
individual follicles, very irregular in shape, are 50-30 p. in 
depth and 30w-in width, Linst^ow's measurements being 65 x 47 ^ 
and Kiessling's 6 ja in the larva. Concerning the shell-gland Lin- 
stow said: "Die Schalendruse ist ein .088-0 .105mm. grosses Organ, 
da3 dorsal von der Mitte de3 einen Keimstockf lugels an der Ent- 
sprechenden Aussenwand der Uteru3 liegt; die Zellen, deren kleiner 
Kern sich intei^iv farbt, sind 0. 0039mm. gross." In the sections 
made by the writer it was found to be a quite irregular structure, 
composed of greatly elongated club-shaped cell 3 with necks of 
different lengths v/hich unite with the oviduct in a region only 



43 



about 30ju». in length and situated just beyond the point of recep- 
tion of the common vitelline duct (Fig. 4). These cells are so 
loosely arranged and their proximal attenuated portions of such a 
filiform nature that they are very easily overlooked, especially 
3ince they are scattered thruout the whole of the dorsoventral 
diameter of the medulla of the region and are interwoven among the 
oviduct, the receptaculum, the vitelline duct and the beginning of 
the uterus. They form by no means such a compact organ as 
Linstow' s description and figure would indicate. The distal ends 
of the cells are about 15 x 10 Jjl in size, while their nuclei are 
about 4^ in diameter. Kie3sling described the shell gland as 
similar to that of S ch. solidus and as follows: "Die Schalendruse 
besteht aus Drusenzellen, welche an feinen Stieichen befindliche 
BlEschen an der Oberflache einer Ealbkugel angeordnet sind and ihre 
Stieichen als Radien nach dexn Mittelpunkte der Kugel senden"; and 
his figures of such a compact region are likewise quite different 
from conditions described here. The uterus forms a mass of coils, 
0.5-0. 6mm. in diameter in the median line, from which a straight 
portion passes ventrally thru the musculature and cortex to the 
opening which is about 20y. in diameter (35 , Linstow) . The 
diameter of the duct is 60^ in the median frontal plane but only 
half that amount a3 it passes thru the longitudinal muscles. The 
measurements of the eggs are according to Eiessling and Linstow, 
respectively, 49 x 34^ and 65 x 43 jji, : they were found by the 
writer to be 53-54 x 31-33^ in sections. 

Our knowledge of the life-history of the species is con- 
fined chiefly to the works of Duchamp (1876 ), Donnadieu (1877) 



43 

and Riehm (1883) who firmly established the well known fact that 
the larva present in the abdominal cavities of various species of 
teleosts develops rapidly in the intestines of fish-eating birds. 
The production of egg3 begins after about 36 hours, while the 
adults live for from three to four days only in the definitjve 
hosts. But apart from these and other closely related details 
which were brought out by Donnadieu. "by means of well conducted and 
controlled experiments, nothing is known, so far as the writer is 
aware, of the development of the oncosphere in the intermediate 
host up to the time when they become distinguishable as small 
larvae. The measurements of the smallest larva found by the 
writer have been given above; Figs. 5 and 6 are of two slightly 
larger specimens, the latter being 6.1mm. in length by 1.34mm. 
maximum breadth. 

Altho the above description shows many discrepancies be- 
tween the species as here dealt with and the European form, the 
writer doe3 not feel justified in separating the two specifically, 
especially in the absence of European material for comparison. 
The thickness of the cuticula, and subcuticula, the dimensions of 
the testes, seminal vesicle and cirrus-sac and the diameter of the 
vagina show the greatest differences, apart from the probable 
confusion by Linstow of the oocapt and receptaculum seminis, while 
the measurements of the eggs as here given are somewhat inter- 
mediate between those by Kieesling and Linstow. But the fact 
that the data given by the latter are apparently the only adequate 
one3 for the adult and that there are not a few discrepancies be- 
tween Kiessling's and Linstow' s accounts restrains the writer from 



44 

looking upon this, the American form, as new. In dealing with 
this question of identity we must also remember that not only- 
does the species vary so much that, as pointed out above, a great 
deal of confusion exists in the earlier literature but that the 
number of host species of the larva as well as of the adult is 
very large as compared to other species of bothriocephalids, hence 
introducing greater factors for variation; and above all that the 
geographical distribution of the wading and diving bird3 harboring 
the mature worms is such that here in America we have many of the 
same species as well as the same genera that occur in Europe. As 
the above record of hosts indicates, the species ce tainly ranges 
widely over Europe and Northern Asia, so that it would be quite 
surprising if it did not occur here in North America, with the 
probable region of transition in Iceland and Greenland on the east 
and Northeastern Siberia and Yukon on the west. However, apart 
from Leidy's and Linton's records it has apparently not been re- 
ported up to the present. 

The material studied by the writer consisted of the follow- 
ing lot of larvae: Nos. 4706 and 4708 of the Collection of the 
United States National Museum; Ch 18a, 16.411, 16.413, 16.414, 
16.419, 17.31 and 17.32 of the Collection of the University of 
Illinois, under the care of Professor H. B. Ward; Nos. II, III, 
IV, and V from the Collection of Mr. H. R. Hill made at Turtle 
Lake, 'Michigan; and Nos. 47, 54, 150, 158, 159, 160, 189, and 190 
of the writer's collection; and the adults contained in Nos. La- 
156, 17.184, and 17.185, C. U. 111. respectively from the intes- 
tines of Merganse r sp . , Podil ym bus podic eps and Coly mbus Kolboellii . 



45 

Genus 2. Schist ocephalus Creplin, 182 9. 

Taenia (part.) Auctorura. 

Hirudo (part.) Linnaeus , 1745. 

Fasciola (part.) Linnaeus, 1767. 

Rhvtis (part.) Zeder,1800. 

iial ysis (part.) Zeder,1800. 

Bot h rlQ C. e .gha jus (part. )Rudolphi, 1808. 

Schistoce phalus(part . ) Creplin, 1829. 
Generic diagnosis; Bothria and external segmentation al- 
readyjdeve loped in the larva. The tip of the scolex retractile. 
Segmentation complete and corresponding to the internal structure 
of the animal. Longitudinal and transverse muscles arranged in 
several alternating aayers (three transverse layers enclosing two 
longitudinal layers). 

Type (and only) species: Sch . solidus (0 .F.MuellerJj. 





Schist ocephalus solidus 


(0. F. Mueller, 1776) . 






(Figs. 7 - 


9.) 






Larval 


stage: 








1734 


Taenia 


Frisch 


1734 


: 395. 


1745 


Hirudo depressa al,ba 


Linnaeus 


1745 


: 250. 


1758 


Fasciola hepatica 


Linnaeus 


1758 


:648 


1761 


Taenia la ta 


Pallas 


1761 


: 410. 


1767 


Fasciola hepatica 


Linnaeus 


1767 


; 1077. 


1776 


Taenia solida 


Mueller 


1736 


: 219. 


1780 


Taenia sasterostei 


Mueller 


1780 


: 22. 


1780 


Taenia sasterostei 


Fabric ius 


1780 


: 320. 


1781 


Taenia acutissima 


Pallas 


1781 


: 76,78. 













4b 


1786 


Taenia *iasterostei 


Batsch 


1786 




2ki4. 


1788 


Taenia BALiriLa 


Schrank 


1788 


* 


49. 


17 90 


Taenia solida 


Gmelin 


17 90 


• 


3079. 


17 90 


Taenia sasterostei 


Abildgaard 


1790 




49-58. 


1800 


RhyJLis spUda 


Zeder 


1800 




297. 


1810 


Bothriocephalus solidus 


Rudolphi 


1810 




57. 


1819 


Bothriocephalus finlirhis 


Rudolphi 


1819 




139,477. 


1819 


Rnt/q-ri nr.Prthffl lllfi f^ril i /lus 


LBuckart 


1819 


* 


46. 


1824 


Bothriocephalus solidus 


N itzsch 


1824 


• 


97. 


182 9 




Baer 


1829 




388. 


? 1863 


^fil-^ fit nee plains rVivnrh- 

ichthydid 


Diesing 


1863 




233. 


1896 


Sch. dimornhus 


Zschokke 


1896 


: 


773. 


1896 


Schistorhvnchus dimor- 
phus 


Zschokke 


1896 




776. 


1897 


Schistocephalus diraor- 


Linton 


1897 




427. 


1898 


Schistoc. solidus 


Conn 


1898 




126. 


1898 


Schistoc. solidus 


Muehling 


1898 




33. 


1899 


Schistoc. solidus 


Luehe 


1899 




52. 


1909 


Schistoc. solidus 


Scott 


1909 




80. 


Adult 


stage : 










1783 


Taenia lanceolata nodosa 


Bloch 


1782 




10. 


1786 


Taenia lanceolata var. 


Batsch 


1786 




167. 


1788 


Taenia nodularis 


Schrank 


1788 


5 


39. 


17 90 


Taenia lanceolata nodosa 


Gmelin 


1790 




307 5. 


17 90 


Taenia sasterostei 


Abildgaard 


1790 




49-58. 


17 93 


Taenia lanceolata nodosa 


Rudolphi 


17 93 




41. 


1800 


Halvsis alnceolato- 
nodosa 


Zeder 


1800 


; 


340. 

















*x ( 


mo 


Bothrioce 


ohalus nodosus 


Rudolphi 


1810 




54. 




1819 


Bothriocephalus nodosus 


Rudolphi 


1819 




140. 




1819 


Bothriocephalus nodosus 


Leuckart 


1819 


: 


58. 




1824 


Bothrioce 


phalus nodosus 


Nitzsch 


1824 




97. 






Schistoce 
phus 


phalus dimor- 




I09Q 




yo . 




1839 


.Schistoc . 


d,imorphus 


Creplin 


183 9 


• 


2 96. 




1845 


Schist oc . 


dimorphus 


Dujardin 


1845 




622. 




1850 


Schist oc . 


dimorphus 


Diesing 


1850 




584. 




1853 


Schistoc. 


dimorphus 


Baird 


1853 


• 


92. 




1854 


Schistoc. 


diraorDhus 


Diesing 


1854 




19. 




1858 


Schistoc . 


solidus 


R. Leuckart 


1858 




129. 




1859 


Schistoc. 


solidus 


Steenstrup 


1859 




475. 




1863 


Schistoc . 


dimorphus 


Diesing 


1863 


• 


232. 




1869 


Schistoc . 


dimorphus 


Willemoes- 
Suhra 


1869 




469. 




1877 


Dibothrium ligula 


Donnadieu 


1877: 




495. 




1881 


Schistoc. 


dimort)hus 


Monniez 


1881 


5 


175. 




1882 


Schistoc. 


dimorphus 


Kiessling 


1882 








1889 


Schistoc. 


solidus 


Loennberg 


1889 


: 


40. 




1890 


Schistoc. 


dAnjqrphus 


Loennberg 


1890 


: 


18. 




1893 


Schistoc . 


dimorphus 


Oisson 


1893 


■ 


15. 




1896 


Schistoc . 


diraorohus 


Ariola 


1896 


* 


280. 




1896 


Bothriocenhalus 

zschokkei 


Fuhrmann 


1896. 








1898 


Schistoc. 


zschokkei 


Puhrraann 


1898 


: 


144. 




1898 


Schistoc. 


so lidus 


Muehling 


1898 


5 


33. 




1899 


Schistoc. 


nodosms 


Luehe 


1899 




52. 




1900 


Schistoc. 


dimorphus 


Ariola 


1900 


:426 





48 



1910 Schist oc. gasterostel Luehe 1910 : 19. 

1911 Schistoc . dimorphus Solowiow 1911 ; 123 



Specific diagnosis: With the characters of the genus. Me- 
dium sized worms, length 30 to 300mm. , breadth 3 to 9. First seg- 
ment or"scolex" 0.4 to 0.8mm. in length and 1.0 to 1.3 in width. 
Strobixa ovate-lanceolate and depressed, maximum breadth anfeferior to 
the middle; hindermost segments narrower and flatter, 0.25 to 1.0 
mm. in length by 1 to 3 mm. in width, forming an appendage up to 
10mm. in length; medium segments 0*1 to 0.5mm. long, posterior bor- 
ders prominent. Shallow median groove on the ventral surface. 

Cuticula 15 to 20,^ in thickness; subcuticula 40 to 65<^. Lay- 
er of internal longitudiaal muscles 15 to 50p-in thickness, herve 
strands 30 to 75|x in diameter. 25 to 30 excretory vessels in tran- 
sections. 

Genital cloaca median, shallow , with a diameter of 90|j- ; no her- 
maphroditic duct. Opening of vagina close behind that of cirrus 
and to one side but not so far as that of the uterus, both alter- 
nating irregularly from side to side. 

Testes extena from the median genital ducts laterally to the 
edges of the medulla, unbroken from proglottis to proglot t is , close- 
ljr crowded, 240 to 480 in number for each proglottis, 85 to 100^ in 
depth, 40 to 65 in width and 55 to 85 in length. Vas deferens me- 
dian, dorsal, closely applied to the seminal vesicle, the whole mass 
0.30ram. in diameter, the duct itself 35 to 60^. Seminal vesicle 
175 x 150(>- , walls 25 ^ in thickness. Cirrus-sac oval in shape, 
immediately below the seminal vesicle, 0.185 - 0.203 x 0.203-0.212 
x 0. 16t)-0» 185mm. in dimensions. &o inner seminal vesicle. Cirrus 



49 

proper not sharply Lpparated from the ductus e jaculatorius ; whole 
surrounded by numerous retractor muscles. 

Vagina 45 to 60^ in diameter just within the medulla. Recep- 
taculum seminis large, 92-104^ in diameter. Sperraiduct unites v ith 
the oviduct close to the ventral wall of the medulla. Ovary with 
large wings consisting of closely arranged tubules, whole organ 
0.6ram. in width, v, ingsO. 10 in length. Ova 15 fA in diameter, their 
nuclei by. . QQcapt 35 to 40f*.in diameter, oviduct 25 to 30^ . Vi- 
telline gland unbroken at margins of the proglottis, from proglottis 
to proglottis and medially, excepting for small areas above and be- 
low the proximal reproductive ducts; individual follicles 55-90 x 
18-26 ^ . OOtype 16-20}*. in diameter. Shell-gland slightly to one 
side of median line. Uterus 85-i35fv in diameter at its middle; 
the terminal portion directed dorsoventrally and lined with cuticu* 
la dis tally; opening at the bottom of a slight invagination of the 
ventral body wall, formed by the rupture of a preexisting cuti- 
cular membrane. 

Eggs, 38-65 x 22-38^ . 

Habitat: As larvae in the body-cavities and coccasionally in 
the stomach anc intestine of bony fishes; adults in the intestines 
of wading and diving birds. 

Authorit y 

Diesing 1850:584. 
Fabriciusl780:320. 
Rudolphi 1810:58. 
»• 1819:140. 



Host Localit y Collector 

Larval stage: 

Gasterosteus aculeatus Frisch 

■ " Greenland Fabric ius 
" " - Rudolphi 

■ " Gryphswald & " 

Berlin 











50 


? Gasterosteus aculeatus 


Baer 


Diesing 


1850:584. 


y » it 




Creplin 


ii 


it ii 


N II 


Bracciano, Italy Parona 


Parona 


1899:8. 


It II 


Rome 


Vinciguerra 


it 


it it 


II II 


Loch Loriston, 
Cove, Scotland 


H.C.William- 
son 


Scott 


1909:80. 


H nunsitius 




Frisch 


Diesing 


1850:584. 


7 it it 




Baer 


it 


it tt 


^ it ti 




Creplin 


it 


it ti 


Cottus scornio 




Zoega 


Rudolphi 


1810:58. 


" poeciloDUS 


L.Stors jon, 
Sweden 


Olsson 


Olsson 


1895:15. 


" bairdii 


Swan R. ,Mont . 


Everraan 


Linton 


1897:427. 


Salmo salar 




Mueller 


Rudolphi 


1810:58. 


it it 


Gryphsv.ald 


Rudolphi 


ii 


1819:140 


ii it 


Basel 


Zschokke 


Zschokke 


1896:776 


Totanus calidrus 




Creplin 


Diesing 


1850:584 


Fulica atra 




it 


it 


it it 


Phoca vitulina 


Gryphswald 


Rudolphi 


Rudolphi 


1819:140 


Rhvnchichthvs 
son sronovii 


Hayti 


We inland 


Diesing 


1850:585 


Rana esculenta 






Luehe 


1910:19 






Gasterosteus bisni- 
nosus atkinsii 


Chamc ook L. , 
New Brunswick 


Cooper 


Cooper 
(the pres 


ent paper) 


Uranidea forraosa 


Port Credit, 
Ontario. 


it 


it 




Adult stage: 










Corvus corax 




Schilling 


Diesing 


1850:584 


■ cqrnix 


East Prussia 


Braun 


Muehling 


1898:34 


Recurvirostra 
avocetta 




Schilling 


Diesinfe 


1850:584 











Ol 


Ardea 




Braun 


Diesing 


1850:584 


ii 




Abildgaard 


hudolphi 


1810:54 


ii 


steixaris Oenf 


Puhrmann 


Fuhrmann 


1896:546 




Schilling 


Diesing 


1850:584 


ii 


alba East Prussia 


Braun 


Muehling 


1898:34 


ii 


ciconia 




Luehe 


1900:19 








Sterna 


hirundo Grvphswald 


Rudolphi 


Rudolphi 


1819:140 


ii 


arctica 


Schilling 


Diesing 


1850:585 


it 


iU^ra - 


ti 


it 


ti it 


M 


macroura Grvphswald 


MUS .2.001. 

Gryphswald 


ii 


ii it 

- 


II 


mi nut a " 


ii 


ii 


rt ii 


Colvmbus spntentrio- " 
palis 


Rudolphi 


Rudolphi 


1819:140 


it 


ti 


Creplin 


Diesing 


1850:585 


ii 


" Pillau 


i.iuehling 


Muehling 


1898:35 


it 


■ Firenze, Italy 


Condoreili 


Parona 


1899:8 


it 


cristatius Grvohsv.ald 


Rudolphi 


Rudolphil81819: 140 


ii 


^lacilis -- 


Abildgaard 


Diesing 


1850:585 


i» 




it 


Rudolphi 


1810:54 


ii 




it 


ii 


» ii 


ii 




Schilling 


Diesing 


1850:585 


ii 


grisei.^ena 




Luehe 


1910:19 


P.Qdice 




Rudolphi 


Dieeing 


1850:585 


n 


* V Pillau 


wiuehxing 


Muehling 


1898:34 


tt 


rubricollis 


Nitzsch 


Mtzsch 


1824:98 


it 


niericollis Bracciano. 

Italy 


Parona 


Parona 


1899:8 


if 


ii _ 


Solov, iov: 


Solow iow 


1911:123 











52 


Larus caoistranus 




Schilling 


Diesing 


1850:585 


" ridibundus 




Siebold,Coll, 
Brit .Mus. 


Baird 


1853:92 


it » 


Rossitten 


Muehling 


Muehling 


1898:34 


" marinus 


Pillau 


it 


ii 


ii ii 


" argentatus 






Luehe 


1910:19 






it it 




Siebold 


Moniez 


1881:175 


Anas £lacialis 




Creplin 


Diesing 


1850:585 


Mersus albellus 




Bloch 


ii 


ii it 


H it 




Schilling 


ii 


ii it 


" merganser 




Bloch 


•t 


ii it 


»t ti 




Schilling 


M 


it it 


it ti 


Pillau 


muehling 


Muehling 


1898:33 


■ serrator 




Abildgaard 


hud ol phi 


1810:54 


H II 




Creplin 


Diesing 


1850:585 


II It 


? Glasvaer, 
Norway 


Loennberg 


Loennberg 


1890:18 


II II 


L.Stors j on, 

J erat land, Sweden 


01s son 


Olsson 


1893:15 


It II 


? Pillau 


Muehling 


Muehling 


1898:34 






Abildgaara 


Diesing 


1850:585 






Schilling 


ii 


ii ii 


Alca pica 




ii 


ii 


it ti 


M t orda 


Leipzig 


C.W. Stiles 


Stiles and 
Hassall 


1894:322 


Totanus calidrus 


Jaederen,N or- 

v:ay 


Loennberg 


Loennberg 


1890:18 


Harelda glacialis 


Pillau 


Muehling 


Muehling 


1898:34 


Fuli.^ula mar i la 


H 


ii 


ii 


ti ii 


Maematoous ostrea 


ii 


ii 


ii 


ii it 



53 

funics atra Portoi'erra j o, Damiani Parona 1899:7 
Id. Elba 

Puffinms kuhli " " " 

Urinator arct icua Luehe 1910:19 

imber M " 

" stellatus " 

Stercorarius para - 

sit icus 

N yroca mariia — " 

hy emails - " H " 

T.nr^ nri v^es cucul - Lincoln, Nebr. H.B.Ward Cooper 

latus (the present paper) 



54 

As indicated in the above synonymy this species was known 
for almost a century, at first as the larval form only and then as 
both larval and adult forms, before it was discovered that the two 
species recognized from the time of Bloch (1783) were one and the 
same. Abildgaard (1790), who called the work T. gasterost ei , 
seems to have been the first to consider the larval form found 
chiefly in stickle-backs to be the same as that found in fish-eating 
birds, since on feeding stickle-backs infected with the larvae to 
geese he obtained the admit form from the intestines of the latter. 
Yet Rudolphi (1810) did not agree with his conclusions but still 
considered that there were two distinct species, namely, Bothrio- 
oephalus nodos u8_ (adult) and B. solidus (larva). And this con- 
tinued until Creplin (1839) united both in one species under a new 
genus, Schistoce phalus dimorph us . Die sing (1863: 233) made a new 
species out of the Sohi s tocephalus found by Weinland (1859) in the 
Island of Hayti in Rhyjphi chthys granov i i , but later writers have 
considered that in all probability it was only the well know 
larval form of this species. Willemoes-Suhm (1869) was evidently 
the first to study the development of the fertilized ovum which 
was later gone into more thoroughly by Schauinsland (1885 : 555). 
Donnadieu (1877) to whom all go back in their comside rations of the 
larval development of Ligula, unfortunately fell into the error of 
considering Schis tocephalus and Ligu la to be not only the same 
specifically but generically. The anatomy was first studied by 
Moniez (1881 : 175), more thoroughly by Kiessling (1883) and still 
later by Fuhrmann (1896) ( under B. zfichok kei sp.nov.) and Solewiow 
(1911). Linton (1897 : 427) is the only one, apart from 



55 

Weinland's record which the writer was not able to locate, who 
has reported the species from America. 

'As regards the correct name of the species it should be 
noted that, altho Luehe (1899 : 52) called the "typical and only 
species" of the ?enue Sch . nodos us (Rud.) and the larval stage 
Sch. solidus (0. F. Mueller), he reverted in 1910 to " Schiat . 
gasterostei (Fabr.) ( = Sch . dimorphus Crepl . ) t! without, however, 
discussing the change. But according to the Rules of Nomenclature, 
Art. 27 (b), the earliest name of the larval stage must hold, so 
that, since Luehe himself considered this to be Sch . solid us 
(0. F. Mueller), the writer makes use of the latter in the present 
paper . 

According to Luehe (1910 : 19) Sch. solidus ranges in 
length from 30 to 300mm. while the maximum breadth varies from 
about 3 to 9mm. and is located ahead of the middle of the strobila. 
As shown in the table below the largest and only sexually mature 
specimen of the 3ix studied by the writer (vide infra) was only 
29mm. in length by 6mm, in breadth. The scalex (Fig. 7) is, as 
indicated in the above diagnosis of the subfamily, not separated 
from the first segment into which it runs insensibly, the whole 
"head" being thus triangular in shape. The bothria are merely 
short median grooves which unit at the very tip not only with each 
other but with a frontal median groove which passes laterally into 
3light emarginations of the edges of the segments. Ifnile these 
emarginations were seen by the writer to be present in the anterior 
segments, gradually disappearing towards the middle of the worm, 
no such "flat leaf like flaps (bothria) on the lateral margins, 



56 

separated from each other on flat surface by a broad, shallow 
sulcus," as described by Linton (1897 : 438) and 3hown in his 
Fig. 4, PI. XXVIII, for the first segment were met with, but the 
posterior border was quite entire, altho, as seen in Fig. 7, not 
very prominent in the vicinity of the median line in adults as 
well as in larvae. The bothria of the mature specimen (Efi of 
the table below) were not present, but the region where they would 
otherwise be was seen to be quite smooth, only a shallow, median 
frontal groove appearing. The whole strobila is ovate-laceelate , 
considerably depressed and provided in the adult with a very 
shallow median groove on the dorsal surface (Fig. 9) which seems 
to be due to the slight protrusion of the median reproductive 
organs, chiefly the cirrus-sacs and seminal vesicles, towards the 
ventral surface (Fig. 8) and the consequent dragging downward of 
the dorsal median tissues. Concerning this matter Linton said 
that "Si. dimorphus is described as having in the larval state a 
longitudinal median furrow on each face. These specimens do not 
exhibit this character; neither do they have anything that can be 
properly called a costa dividing the two bothria." While in the 
specimens studied the dorsal groove was present not only in the 
adult but (not so well marked) in the larva, a similar ventral 
groove was also noticed in sections of the anterior end of one of 
the latter. Both grooves, however, are in any case so shallow as 
to be easily overlooked in alcoholic specimens; they seem to be 
of only secondary importance since they are apparently quite vari- 
able in their nature. While the segments in the anterior region 
of the strobila are very broad and comparatively thick, short, and 



57 



from 0.1 to 0.5mm. in length, posteriorly the strobila is consider- 
ably smaller and flatter, especially in mature individuals, but 
even in larvae the segments being much more irregular in outline 
and as much as 1mm. long (0.75 in the only ripe specimen studied). 
The segmented condition of the strobila, in contrast with that of 
Li^ula, is rendered more apparent by the prominent posterior bor- 
ders of the anterior and middle proglottides which at the margins 
produce the characteristic saw-tooth effect. The following table 
gives the measurements of two specimens with those by Linton for 
comparison: 



Number 


72 


H.7 


4737, U.S.N.M. 


Length 


17mm. 


39mm, 


33mm . 


Max. breadth 


5.5 


6 


6 


Length of "Cauda" 


1.64 


10 


? 


Breadth of same 


1.1 


3-3 


3.5 


Med. segs., length 


0.16 


0.37-0.46 


0,35 


Post. 3egs., " 


0.35-0.40 


0.40-0.75 


? 


First 3eg., " 


0.46 


0.46 


0.80 


Breadth anteriorly 


0.48 


0.46 


0.80 


" posteriorly 


1.11 


1.11 


1.30 


Length of bothrium 


0.074 


Absent 


? 


Condition 


Larval 


Adult 


Larval 



Since the essential features of the internal anatomy of 
this species have been worked out by the European workers, only 
the striking similarities and differences to and from the data 
given in particular by Kiessling, Fuhrmann and Solowiow will here 
be dealth with in support of the writer's contention, in the ab- 



_ 53 

sence of European material for comparison, that here in America 
we have the same species as that found in Europe. And it will 
be considered that, as brought out by Luehe in his three contro- 
versial references (1897b and 1899a : 715) and by Cohn (1898 : 
126, footnote), S. zschok kei Fuhrmann 1898 is synonymous with 
S. solidu s, since many of the data given below will be seen to 
compare more favorably with those published by Fuhrmann than with 
those by either Kiessling or Solowiow. 

According to Kiessling thecuticula is from 15 to 18^ in 
thickness and divisible into two layers, of which the inner and 
lighter is from 8 to 9 ^ thick, while the outer is striated or 
granular. Fuhrmann described a cuticula only 7pL in thickness 
and divided into two layers, and Solowiow gave the thickness of 
the "homogeneous cuticula" as 23^ , but Minckert (1905a : 402) 
said that the comidian or pseudociliated layer, present in many 
bothriocephalids, was quite evident in S. nodosus but absent on 
the posterior borders of the proglottides. Here the cuticula was 
found to be 15^ in thickness, excepting on the posterior borders 
where it was only 5^ , and to be divisible into two layers, the 
outer of which, a little thinner than the inner, was much lighter, 
granular in consistency or somewhat striated with, however, a more 
or less uniform external boundary. It seems to be easily 
separated from the inner stratum, the bounding line, in reality 
the innermost portion of the external layer, being in most places 
very light. In fact the brightness of this inner layer of the 
outer stratum indicates the degree of separation of the two layers 
in the process of sloughing off the outer, which can be easily 



59 



followed in sections aa described by Kiessling. But this de- 
scription applies only to theafiult stage, for in larvae the cuti- 
cula, altho of the same thickness, shows an outer decidedly pseu- 
dociliated layer only 4 ^- in depth. The subcuticula, 88.5 ^ in 
thickness in the median line according to Solowiojn, was found to 
be from 40 to about 65 ^ , Kiessling having given the limits as 
from 39 to 38^ . While the parenchyma is, a3 described by the 
authors, very fine meshed, calcareous bodies are present in com- 
paratively small numbers, particularly just beneath the subcuti- 
cula of the larva. Their maximum dimensions were found to be 
23 x 13|l. 

The musculature has been well described by Kiessling and 
Fuhrmann, so that it needs only to be said that in sections of 
mature proglottides the writer found that the outermost layer of 
transverse muscles as well as the outer longitudinal layer were 
much less numerous and hence well defined than in the larva. 
Whereas Kiessling gave the thickness of the external and internal 
longitudinal groups, which on account of their compact nature were 
found to be more uniform in thickness than the trans vers layers, 
as 8-33 and 16-49 ^ , respectively, and Fuhrmann as 4 and 8 p. , 
the writer found them to be 17 and 30-40^ . 

The nerv.ous system was first studied In detail by Niemiec 
(1888 : 9) and later more thoroughly by Cohn (1898 : 136) who 
summarized its structure in the following words: "Von dem vordersten 
Theil, den Ganglien una der Commissur, Ziehen die Haupt strange und 
13 Nebennerven ruckwarts (six associated with each chief strand). 
Die Nebennerven theilen sich dichotomisch in zwei Ebenen, der 



30 

frontalen und radiaren, ein Theil des Theilfasem ruckt zwischen 
aussere Transversal und Lang smu skein, der andere bleibt weiter nach 
innen zu zuruck, und diese Nerven treten einerseits unter einander 
durch Ringcommissuren, andrerseits durch radi'are Fasern mit den 
Hauptnerven in Verbindung. " Whereas Kiessling gave the diameter 
of the chief nerve strands as 38 |n and Sidjyiow as 67. , here 
they were found to be from 30 in mature proglottides, to 75^ in 
the anterior segments. The ganglia were found to have a diameter 
of from 55-85 , compared with 77 by Kiessling. 

In transections from 25 to 30 excretory vessels were seen 
in the medullary parenchyma with diameters ranging from 29 to 63^. 
Fuhrmann gave 24 as the number while Solowiow gave their size as 
13.9 to 23. 3|*^. Foramina secundaria are to be 3een piercing the 
cuticula here and there, but they are not very numerous. 

As indicated in the diagnosis of the subfamily the repro- 
ductive organs appear close behind the scalex. In one toto pre- 
paration of a larval specimen, No. 72 of the aoove table, the 
earliest traces of their rudiments were present in the 18th pro- 
glottis, or 3.96mm. from the anterior end, while in the only mature 
specimen, H.7, they were in the 16th proglottis, a few eggs appear- 
ing in the uterus of the 17th. The cirrus and vagina were found 
to open close together in a very shallow and sometimes quite obli- 
terated genital cloaca. With a maximum diameter of about 90|jl , 
the vagina behind the cirrus, but very slight either to the right 
or left and not according as the uterine opening further back like- 
wise alternates irregularly but with a greater amplitude. The 
three apertures form almost a right-angled triangle, as described 



62 

"by Kiessling, but, as was pointed out by Luehe (1899a :716) this 
arrangement is by no means constant but varies with the state of 
contraction or relaxation of the whole strobila and hence cannot 
be considered as specific. 

The testes are arranged in a single layer in the dorsal 
portion of the whole of the medulla not only in the larval but also 
in the adult, as described by Fuhrmann, the majority of the 
excretory vessels being situated towards the ventral side of the 
medulla, and are absolutely continuous from proglottis to pro- 
glottis. Their number in transections is from 30 to 40 (30-35, 
Kiessling) and in sagittal from 8 to 12 for each proglottis, thus 
making the total from 240 to 480 or over 300 on the average, which 
stands in distinct contrast with the number of about 100 given by 
Fuhrmann. The latter gave their dimensions as 80 x 34y~, Kiessling 
as 16 to 36 |^ in young and 149 in mature animals, and Solowiow as 
68-93 pL ; while the writer found them to be from 85 to 100 [k in 
depth, 40 to 35 in width and 55 to 85 in diameter in frontal sec- 
tions. They are very closely crowded together in the proglottis. 
The vas deferens form3 a compact mass of coils situated in the 
median line dor sally and slightly posterior to the vesicula semi- 
nalis to which it is closely ap. lied as a sort of cap. While the 
diameter of the whole organ is about' 0.3mm. that of the duct it- 
self is from 35 to 60 Ja- when distended with spermatozoa. Kiessling 
gave its diameter as 38 , and Solowiow as 16.3. The large thick- 
walled seminal vesicle (Fig, 9) situated immediately above the 
cirrus- sac was found to have a maximum depth of 175 yc and transverse 
diameter of 150 ja. , as compared with the 92 ^ of Kiessling and the 



63 

80 y~ of Fuhrmann. Its wall 9 are very muscular, about 25 U in 
greatest thickness and covered both internally and externally with 
numerous nuclei which ar, : ; respectively epithelial and parenchyma- 
tour or myoblast! c in their nature. TJithin the cirrus-sac the 
vas deferens is much coiled but not enlarged to form any secondary 
vesicle nor 3harply separated into an ejaculatory duct and cirrus 
proper. The sac itself is oval in shape, the ventral end being 
the smaller, and the proximal end somewhat invaginated by the 
seminal vesicle. Its size i3 shown in the following table: 

Ki es sling Fuhrmann Sol owiow The wri ter 

Depth 0.347mm. 0.25mm. 0.804mm. .185-0 ,303mm. 

Width 0.192 » 0.13 ■ 0.174 " 0.20-3-0.312 » 

Length ... ... ... 0.166-0.185" 

Its wall, about equal in thickness to that of the seminal vesicle, 
is, however, more open in texture, the Myoblast! c nuclei of the 
obliquely arranged muscle fibres being scattered thruout its dia- 
meter (Fig. 9). It is, furthermore, not sharply separated either 
externally or internally from the surrounding parenchyma and the 
numerous stout retractor muscles of the cirrus, respectively. The 
latter, in fact, constitute practically the whole of the contents 
of the sac apart from the duct itself. The only protruded cirrus 
seen had a length of 70f^, as compared with the 0.3945mm. given 
by Solowiow. 

The vagina, the opening of which is usually situated 
about 50 J* from that of the cirrus at the bottom of the shallow 
genital cloaca, above mentioned, ha3 a diameter of from 45 to 60 |a. 



63 



at the first bend in its course within the meiullary parenchyma. 
Soon after it enters the latter it becomes thin-walled as pointed 
out by Fuhrmann, owing to the thinning out of the cuticula and the 
substitution of the proximal nucleated epithelium for the same, 
altho more peripherally much flattened nuclei are to be 3een be- 
neath the cuticula and crowded close to the basement membrane. 
In other words the gradual replacement from within outwards of the 
cuticula for the original epithelium may be followed very easily 
in the walls of the vagina. The duct gradually enlarges to form a 
much elongated receptaculum seminis (Fig. 8) with a diameter of 
92-104 jtt . (9-31 (a., according to Kiesslingl) and sharply separated 
from the spermiduct, which, however, was not found in the sections 
made to unite with the oviduct close to the dorsal transverse 
musculature as 3tated by Fuhrmann, but close to the ventral wall 
of the medulla. The ovary consists of two large wings (Fig. 8), 
composed of clossly crowdel tubules, lying immediately upon the 
ventral transverse muscles and united by a much smaller isthmus, 
the whole having the width of 0.64mm, a3 compared ?/ith the 0.38mm. 
of Solowiow. The average length and depth of the wings are, 
respectively, 105 and 90^. Ova from the isthmus and more median 
portions of the wings of the ovary were found with a diameter of 
13 while their nuclei were 5 fx. . The respective measurements by 
lies sling and Solowiow were 9 and 6 f- and 13.9-33.3 and 1.5-3 ^ . 
Fuhrmann stated that one of the most important differences between 
his Sen . Zsoh okkei and Sch . solidus_ was the presence in the former 
of an oocapt, but Luehe (1899a : 717) claimed that this structure 
was in all probability overlooked by Kieasling. It arises from the 



34 

posterior aspect of the isthmus almost in the median line with a 
diameter of from 35 to 40 |A- . The oviduct, according to Kie3sling 
has a diameter of 13p~ or to Solowiow of 37f*. ; here it was found to 
be from 25 to 30 ^ between the entrance of the vagina and that of 
the common vitelline duct, which two points are close together as 
in L. intestinalis . The common vitelline duct was found enlarged 
some little distance from its opening into the oviduct to form a 
vitelline reservoir having a diameter of 30 \l (23 fv, Kiessling) . 
The vitelline follicles are extremely numerous and closely crowded 
together in a layer with a maximum thickness of 85^ situated be- 
tween the inner longitudinal and middle transverse muscles (Fig. 8). 
They are continuous at the margins of the proglottis, as they are 
from joint to joint, and broken only in limited elliptical areas 
above and below the reproductive ducts in the median line, as stated 
by Fuhrmann. The size of the individual follicles is according 
to Kie33ling 56-107 x 55^ and to Solowiow 18 x 37y- ; here they 
were found to be 58-87 x 18-36 f , the larger dimensions being 
the dorsoventral diameters. Just beyond the entrance of the common 
vitelline duct the oviduct enlarges to form the ootype with a 
diameter of 16 ^ (20 j^, Kiessling) and surrounded by the shell- 
gland which is situated just above the median frontal plane and 
somewhat lateral. Thruout its course the oviduct is lined with an 
epithelium in which prominent nuclei but no dictinct cell boundar- 
ies were seen and from which numerous cilia protrude into the lumen. 
In the ootype these cilia are much more noticeable. From the oo- 
type the oviduct passes ventrally with a few coils then across the 
median line close aoove the receptaculum semini3 as the beginning 



55 

of the uterus. The latter gradually enlarges as it passes forward 
while crossing the median line several times, until at about the 
middle of it3 course it ha3 a diameter of 85 to 135 . A3 re- 
gards the terminal portion of the tube the writer found that, as 
Fuhrmann observed: "Der Endtheil der Uterus verengert sich and 
verlauft von der Dorsalflache (the median frontal plane in which 
the last transverse coil is situated) direkt ventral, urn regel- 
ma , 33ig abwechselnd links oder rechts neben der Vagina auszumunden" 
(vide supra). 

Sections show that the actual opening is formed by the 
rupture of the bottom of a cup-like invagination of the cuticula 
from the ventral surface, which meets the end of the duct with a 
diameter of from 25 to 40^ . As Fuhrmann stated, "Dieser Aus- 
fuhrgang der Uterus ist von der Stelle an, wo er ins Rindenparen- 
chym tritt, wie die Vagina und der Cirrusbeutel , von zahlreichen 
Parenthymmuskeln umhullt Und von einer der Korper "^cuticula ahniich- 
en membran ausgekleidet , ■ but the cuticula was found as 3uch only 
near the opening, since only halfway back along tnia dorsoventral 
limb of the organ flattened nuclei could be distinctly seen. In 
other words the flattened epithelium of the uterus, which showing 
only a few scattered nuclei was described by Kie33ling as a "fine, 
structureless but elastic membrane" passe 3 insensibly into the 
cuticula near the opening, no distinct line of junction between the 
two being discernible, which latter i3 also applicable to the 
similar structure of the vagina. 

The dimensions of the ellipsoidal eggs in the 3ctions of 
the uterus were found to be 62-65 x 33-35 ^ . Kie3sling gave them 



66 

as 49 x 37^ and Fuhrmann as 70 x 29 ^ , but in discus 3ing the 
latter Luehe (1899a : 718) remarked that not only did he find 
variations from 38 x 33 to 56 x 38 ^ in the size of the eggs in 
■Mtterial of B. gachokk ei sent to him by Fuhrmann, but that in 
general even greater variations than these are to be found in 
other species -according to the various writers. 

Our knowledge of the life-history of this species dates 
from the time of Abildgaard (1790) who, wa3 mentioned above, was 
the first to experiment with the larval individuals found in fishes, 
Creplin (1829) united the two forms which were considered to be 
two separate species into one species, evidently on the basis of 
the previous work, especially Abildgaard 'a (cf . Donnadiea, 1877f : 
340-341), while Donnadiett in his elaborate experiments on the life 
history of Ligula unfortunately did not differentiate between it 
and Schi stocephalus . The development of the fertilized embryo 
into the oncosphere was first studied by Wiilemoes-Suhm (1869) and 
later more in detail by Schaumsland (1835 : 555), since when nothing 
of special importance has been added so far as the writer is aware. 
Hence up to the present we know no details of the development of 
the oncosphere into the larva in the intermediate host, as is the 
case with most of the bothriocephaliis . 

As regards the identity of the material studied with the 
European species it will be seen from the above comparisons that, 
while there are many discrepancies among the data given by Kiess- 
ling, Fuhrmann and Solowiow, those by the latter departing the 
farthest in many respects, the resemblances so outweigh the dif- 
ferences as to make the erection of a new species unjustifiable. 



67 

The thickness of the cuticula, the diamter of the excretory vessels, 
the dimensions of the seminal vesicle, the ovary and the eggs, 
which constitute the majority of the differences, might easily be 
explained by differences in age of the material studied, but the 
number of testes (100) as given by Fuhrmann can scarcely be recon- 
ciled with that as given by the writer (300+), altho his dimen- 
sions of the organs agree with those given here perhaps better than 
do t;.ose by Kie3sling or Solowiow. On the other hand we must bear 
in mind other facts which doubtless in the long run are more im- 
portant than a comparison of the details of tha anatomy of this 
evidently highly variable species, namely, the geographical distri- 
bution of the hosts. Altho we cannot place so much emphasis on 
Fabricius' finding T. gast ero ster in the type larval host as long 
ag.vO as 1780 in Greenland, we must remember that here in America 
there are, a3 in the Cf.se of L, inte s tinalis ? not only a number of 
the s:,me genera but also of the same species of the larval as well 
as the adult host as in Europe, so that from this alone we would be 
justified in expecting to find the same species of Schi s tocephalus , 
especially since it infests such a number of different host 
species. But, on the other hand, it is a very surprising fact 
that apart from Linton's report of the larva from Montana evidently 
no one has up to the present found the form in any of the numerous 
fish-eating birds of the continent. 

This evident infrequent occurrence of the. species is illus- 
trated by the fact that the material used for study by the writer 
consisted of only four lots: Nop. Sib and 72 from the body cavities 
of Uranldea formosa , taken from the stomach of Lota m acul osa, and 



68 



190 from the coeiome of GaateroBteaa Mapino aua atkinall , of the 
writer's collection; and No. 17.133 of the C. U. 111. from the 
intestine of Lophodytes cucullatus, the only mature specimen avail- 
able, — • in all only six specimens. 

Subfamily 2. HAPL9B0THRI INAE subfam. nov. 

Strobila formed by the subdivision of the segments of 
a primary strobila. Scolex of the latter a cylindrical, somewhat 
clubshaped organ bearing four ever3ible proboacidea similar in 
structure to those of the Trypanorhyncha; scolex of the former or 
secondary (definitive) strobila merely the slightly modified fore- 
most segment, provided with shallow dorsoventral depressions ana- 
logous to the bothria of other bothriocephalids . An elongated 
neck may be said to be present in the primary strobila. Segmen- 
tation of the primary strobila resulting in the formation some 
distance behind the soolex of a comparatively small number of long 
narrow segments which in turn subdivided anteriorly to form the 
segments of the secondary or. final strobila. Segmentation in the 
latter thus beginning immediately behind the secondary 3colex, but 
complete in it3 anterior region only. Genital organs simple in 
each proglottis. Genital openings surficial, ventral and median 
as in the Diphyllobothriinae . Ovary and shell gland median, re- 
spectively ventral and dorsal. Vitelline follicles in the medul- 
lary parenchyma, as are the testes, both within the nerve trunks. 
Testes separated into two lateral fields by the median excretory 
ye33el and the genital organs in the median line. Vas deferens en- 
larged to form a large non-muscular seminal vesicle before entering 



the cirrus- sac. Cirrus armed with minute spines. Receptaculum 
seminis medium sized, 3harply separated from the spermiduct. 
Uterus divided into a much coiled proximal uterine duct and a large 
uterus-sac, as in the Ptychosothriidae . 

Type genus: Haplobothrium Cooper 

Altho we know a3 yet comparatively little about the life- 
histories of the bothriocephalids, the definitive 3colex and 
strobila develop directly from the larval stage, known as the 
plerocercoid, present in the intermediate host. Thi3 is certainly 
the case with Ligula, Schi s toceph alus , Diphyliobothrium latu m, 
Cyathocephalus truncatus and Iriaenophorus . As a matter of fact 
in all of these the scolex is already more or les3 well formed, 
before thdjlarva reaches the final host, while the plea&cer coid 
continues to grow and 30on snows the beginnings of segmentation 
which marks the young strobila. Consequently the writer feels 
that we must look upon what is called here the primary strobila 
of Haplobothrium as the true 3tro'cila, homologous with the young 
strobila of other bothriocephalids, altho what was formerly con- 
sidered to be the strobila ia quite similar, apart from the absence 
of external segmentation in it3 posterior region, to that of other 
members of the order. Even tho it is provided with a very aber- 
rant scolex region, — and we must remember that the scolex i3 no 
more sharply set off from the rest of the larva in other species, 
such as D. l^-jtxim, — the young and as yet unsegmc-nted primary 
strobila may be considered to be a typical plerocercoid. 

The nerv.ous 3ystem consists of two chief strand3 united 



70 

anteriorly by a commissure, which is doubtless relatively larger 
than in other forms on account of the neighboring proboscides to 
which it sends large branches, as in the larvae of Ligu la and 
Sohistoo epha lus, for instance. Its excretory system is likewise 
ouilt on the typical plan, the posterior connections with the ex- 
terior being, in fact, quite like those of B. scorpi i . On the other 
hand, it can be seen from the description of the development given 
below that the terminations of the nerv".ou3 and excretory system 
in thesecondary strobila, both anterior and posterior, support the 
view that the latter is not homologous with the strobila of other 
bothriocephalids . For what was formerly described as the ring 
Commissure must now be considered as merely a secondary development 
due to the fusion of the severed ends of the chief strands, which 
statement is also applicable to the terminal vesicle of the excre- 
tory system. And this in spite of the fact that the secondary 
scolex is quite similar to the true scolex of other species in that 
it is supplied with two sets of muscles which are not found in the 
foremost segments but are peculiar to the 3colex. Since there is 
considerable evidence in the litefeture of cestodes to show that 
the prominent posterior borders of the foremost segments of many 
species are developed as accessory organs of attachment or for 
locomotion (cf. Spengel, 1905 : 281), we might well ask ourselves 
whether external segmentation in Haplobothrium, particularly since 
it is confined to the anterior region of the secondary strobila, 
is palingenetic or coenogenetic in its nature. The facts that no 
such appendages are present in the primary strobila and that the 
posterior end of the secondary one i3 non- segmented apart from 



71 

the sets of reproductive organs, would seem to point to the 
Original condition being one in which external segmentation was 
absent as in Ligula or Tri&ensphorus . Since, however, in the 
middle region of the secondary strobila there i3 an actual corres- 
pondence between the external and the internal segments, it is 
quite probable that the external segmentation is much older than 
might at first appear, while the ligulate condition of the poster- 
ior end may have developed secondarily. Andjwe must remember, 
too, in thi3 connection that, according to Luehe (1898 : 285) Li- 
gula has descended from fully segmented both riocephalids . 

As regards the remaining characters of the subfamily the 
writer met with not a few difficulties on account of the fact that 
up to the present there is only one genus and one species known. 
Those given are consequently based on a comparison of the species 
with the neighboring subfamilies and in particular with the Diphyl- 
lobothriinae to which it is most closely related. It differs from 
the Diphyllocothriinae , however, in that the genital organs are 
simple in each proglottis; the vitelline follicles are medullary; 
the tests are within the nerve trunks; the seminaljvesicle i3 not 
strongly muscular; the cirru3 is armed with minute spines; the 
receptaculum seminis is medium si^ed; while the uterus is divided 
into uterine duct and uteru3-sac as in the Ptychobothriidae . 



73 

Genus 1. Haplobothrl um Cooper 1914 e.p. 
Haplo b othrium Cooper 1914b : 1-3 
Haplobot hrium Cooper 1314b : 115 

Borders of the terminal disc of the secondary aoolex and 
of the posterior auricular appendage s of both the scolex and 
anterior segments provided with minute spines which disappear with 
the appendages farther back. Nervous system consi3t3 of two chief 
strands situated in the medullary parenchyma outside of the vitel- 
line follicles, uniting inthe anterior end of the secondary strobila 
to form a secondary nerve ring, and eight collateral strands, four 
arranged around each main tract, the latter in the jointed portion 
of the strobila only, but in the true 3colex to form an irregular 
transverse commissure situated among the proboacides. Excretory 
system composed of one large median and slightly dorsal vessel and 
two smaller lateral and ventral, all uniting in the secondary 3colex 
behind the nerve ring to form a vesicle. No genital cloaca, opening 
of vagina close behind that of cirrus, towards the anterior end of 
the proglottis, that of the uterus much farther back. Sphincter 
vaginae present. Vitelline glands in numerous follicles arranged 
cylin Irically around the testes, both continuous from joint to 
joint, leaving clear areas opposite the central genital ducts; large 
vitelline reservoir. Vas deferens provided with a sperm-reservoir 
at its posterior end near the middle of the proglottis; whole 
course of the duct dorsal to -the uterus-sac. Uterus-sac when gra- 
vid occupies the whole of the middle of the proglottis. 

Type species: H. globuli forme Cooper 



73 

The characters here given are in reality those of major 
importance in the species left after what are considered by the 
writer to be subfamily characters are removed. The difficulties 
in defining the genus are great owing to the facts that it 3tands 
alone in the subfamily, contains only one species and is, further- 
more, &e aberrant in many respects. 

Haplobothrium globuli f orme Cooper 1914. 

.,914 Hap 1 obo t hr ium gl obul i f o rme Cooper 1914a : 2 
1914 Ha-olobothrium globulif orme Cooper 1914b : 115. 

Specific diagnosis: With the characters of the genus. 
Small worms, primary strobila up to 70mm. in length, secondary 
to 110mm., with respective maximum breadths of 0.3 and 0.6mm. 
Primary scolex 0.35mm. in diameter, indefinite in length, bulbs 
0.40-0.45 x 0.06 x 0.07mm.; secondary scolex, 0.4-0.5 x 0.35-0. 4mm. 
Auricular appendages disappear at about the 25th segment in normal 
secondary stro'oilas. Foremost secondary segments tetragonal, 
middle and posterior much elongated and considerably depressed. 

Cuticula 3-4^. in thickness, subcuticula 35^. Chief 
nerve strands 18^ in diameter, narrowing intersegmental^ . Ter- 
minal excretory vesicle 20-40^ in diameter. 

Genital organs begin at about the 15th proglottis. Open- 
ings of cirrus and vagina 0.02-0. 07mm. apart. 

Testes spherical to ellipsoidal in shape, 70-115 (a in 
maximum length; SO in each segment. Va3 deferens median, elonga- 
ted, only slightly coiled, 10-55^a in diameter. Vesicula seminftlis 
broadly spindle-shaped, 140 x 90(> . Cirrus 20-30 in diameter; 



74 

cirrus- sac, 0.16-0.21 x 0.14-0.16 x 0.18-0. 20mm. 

Vagina 30-30f* in diameter at its opening, 56 (w. in its 

enlarged distal portion. Receptaculum seminis 30-45yL in diameter, 

o 

gpermiduct 5-lOjc and very muscular. Ovary hi^p,erepiiorm, the 
limb3 being directed posteriorly and often fused with each other, 
the isthmus narrow. Ova from latter 10-13 fx in diameter, their 
nucleu, 7^ . Oocapt 15 to 25 ^ in diameter, oviduct 8-15 1>- . Two 
vitelline ducts, each 6^ in diameter; vitelline reservoir 25 to 
55^ ; follicles spherical to ellipsoidal in shape 8-50 ft in diame- 
ter, very numerous and closely crowded. Ootype 20 y> in diameter; 
shell-gland irregular in shape, poorly developed. Uterine duct 
enlarged proximally with few coils, smaller distally and more 
coiled, median, 25 to 55 y in diameter; uterus-sac elongated, 
filling most of the medulla when granid; uterus opening a small 
median elongated slit, situated near the posterior end of the sac. 
Eggs, 60-70 x 40-43(^ . 

Habitat: In the intestine of the host. 

Host Local ity Coll e ctor Autho rity 

Ami a calva Go-Home Bay Cooper Cooper, 1914b : SI 

(type host) Muskoka, Ontario 

" " Havana, Illinois H. B. Ward Cooper 

(the present paper) 
" " Fairport, Iowa " " 

Type specimen: No. 33.1 in the writer 1 3 collection. 

Co- types : Nos. 33.2 and 33.3 of the same, and in 

the Coll. Univ. 111. 



75 



In a preliminary paper on the systematic position of this 
species the writer (1914*: 1) described the scolex as " ... un- 
armed, although the edges of the terminal disc and auricular appen- 
dages of both scolex and anterior proglottides are provided v/ith 
very minute spines. Bothria, two shallow depressions on the dorsal 
and ventral surfaces, very simple in structure," and in the 
detailed description which followed (1914b) the organ was dealt 
with (p, 83) as follows: "The scolex is quite small, simple exter- 
nally and "with the unaided eye can scarcely be distinguished from 
the first joints. It is shaped roughly like a rectangular solid, 
hollowed out laterally to form simple depressions and dorsoven- 
trally the shallow bothria or organ of attachment. The summit is 
somewhat prolonged as a low pyramidally- shaped disc, quite com- 
parable to that ( "Scheitelplatte") found in the members of the 
subfamily Triaenophorinae Luehe 1899 . . . The opposite end of the 
scolex is modified to form two pairs of auricular appendages 
closely resembling internally as "veil a3 externally those of the 
foremost joints." Furthermore in both papers it was emphasized 
that the scolex differs little in structure apart from the nervous 
and excretory systems from the first segments and that the simple 
bothria, whence the generic name, seem of little functional impor- 
tance as compared to those of other species, while the auricular 
appendages of both scolex and foremost joints with their borders of 
minute cuticular spines probably act as accessory organs of attach- 
ment . 

Since then the latter view has been rendered still more 
highly probable, altho as yet no observations have been made on 



76 

the living worms in their relation to the wall of the host's intes- 
tine, by the discovery that the so-called scolex (Figs. 13 and 14) 
is not in reality the 3C0lex but only a slightly modified anterior 
segment, while the functional scolex is something quite different 
from anything present in the whole order, so far a3 the writer is 
aware . 

As shown in Figs. 10, 11 and 13, the 3colex consists of the 
slightly enlarged anterior end of the original plerocercoid or 
larva from which protrude fou r prob osci des , the whole somewhat 
resembling a hydra or reminding one of the Irypanorhyncha . And, 
as will be presently seen, the latter comparison is a very apt one. 
Fach proboscis consists of a permanently protruded base or stump, 
indicated in Fig. 11, about 85^ in length and 45-55^ in diameter, 
somewhat conical in shape and thickly set with minute backwardly 
directed cuticular spines which pass on to the neighboring portions 
of the end of the larva for a short distance, thru which passes the 
proboscis proper with about the same diameter. The whole forms at 
first sight a continuous tentacle gradually diminishing in size to 
the pointed end. These tentacles attain a length of 0.35mm., in- 
cluding the base, when fully evaginated and are directed almost at 
right angles to the longitudinal axis of the larva, their bjses being 
however, turned slightly forward (Fig. 11). Within the scolex the 
tentacles are accommodated in elongated cylindrical muscular sacs, 
quite comparable in structure to the bulbs of the Trypanorhyncha . 
These lie freely in the loose parenchymatous tissue in the diagonal 
diameters of the region. When the proboscides are invaginated, 
they have a length of 0.45mm. -with a diameter of 0.07, or 0.40 x 



77 

0.06mm. when the tentacles are protruded. The walls of the 
bulb (Fig. 17) are composed to two thick layers of muscles, an 
outer longitudinal or somewhat oblique, much the heavier of the 
two, and an inner circular lined, lined with a cuticula-like 
layer i on the inner border of which numerous flattened nuclei 
appear. The walls are attached to the edge of the 3tump, and 
these layers have the same relative arrangement as that of the 
cuticula and cuticular muscles on the outside of the body, only 
being in the reverse order. Continuous also with the edge of the 
stump are the walls of the proboscis proper, which consist of a 
thin external layer of cuticula and only feeble cuticular muscles. 
Attached to the walls internally thruout its course are the retrac- 
tor muscles of the proboscis which pass backward and become at- 
tached to the posterior end of the bulb. These can be seen best 
in longitudinal sections where the proboscis i3 retracted, for 
then they are closely crowded and much thicker, and their attach- 
ment to the inner end of the proboscis is shown nicely. In the 
retracted condition the latter i$, of course, hollow, the narrow 
cavity, often triradiate in transection (Fig. 17), being easily 
followed into the bulb for about one- third of its length. Closely 
applied to the cuticula of the tip of the proboscis appear in some 
cases gland-like cells taking the counterstain quite like those 
behind the bulbs to be described below. They are shown in Fig. 17. 
Apart from the structures already described, the contents of the 
bulbs and consequently of the proboscides to a certain extent, 
consists of a small amount of loose parenchymatous tissue, and what 
is evidently a good deal of nervous tissue coming into the posterior 



78 



end of the organ (vide infra) . 

Evagination of the probosoidea is obviously brought - about 
by the contraction of the muscles in the walls of the bulbs, but 
the body wall in the vicinity of the latter probably greatly 
assists, since its muscolature ia well developed. Some distance 
behind the posterior ends of the bulbs the latter consists of a 
ring-like layer of locsely arranged main longitudinal fibres 
occupying the middle one-third of the radius of the nearly circular 
cross-section, no transverse fibres, but comparatively strong 
cuticular muscles, of which the inner longitudinal layer is the more 
pronounced. Farther forward this main longitudinal group gradually 
gives off small fibres towards the cuticula a3 they themselves 
diminish in number and 3ize, until at the level of the hinder ends 
of the bulbs only a few of the latter fibres are left just beneath 
the subcuticula, while the outer series has formed a compact layer 
situated close to the longitudinal cuticular fibres (and hence 
outside of the subcuticular nuclei) but separated from them by a 
thin stratum of circular fibres. And this continues to the tip of 
the scolex, mo^t of the remaining inner longitudinal muscles being 
located at the ends of the transverse and dorsoventral diameters of 
the transection. ' > In region of the bulbs the body wall is 
thus quite muscular, and in all probability assists the bulbs in 
evaginating the proboscides by compressing the whole of the paren- 
chyma surrounding them. Between the bulbs and right beneath the 
tip of the scolex a few transverse and sagittal fibres are to be 
found, while just beneath the bases of the 3tumps of the proboscides 
the outer longitudinal muscles unite with the longitudinal cuticular 



79 

fibres in formwj r\ -shaped loo^s surrounding the diagonal quadrants 
of the 3colex which accommodate the bulbs. These loops are evi- 
dently for the control of the direction of the proboscis stumps. 

In working up the original description of this species the 
writer was at a loss to see how the formation of proglottides 
could take place continuously since there was considerable evidence 
to show that only a limited and more or less definite number of 
segments were present. Concerning the anterior and externally 
segmented region of the 3trobila he said: "In many chains this 
region of the strobila is subject to considerable variation. It 
was observed that now and then one of the longest proglottides 
was provided with one or two additional pairs of appendages, 
generally abortive and situated anteriorly 3ome distance apart. 
In a few cases staining and clearing brought out a distinct divi- 
sion of the parenchyma, especially posteriorly, in to what seems 
to be the beginnings of a division of the longer proglottis into 
several smaller ones. Furthermore in one strobila an undivided 
region was intercalated betv/een two jointed regions, the second of 
which was followed by the normal posterior end. Young scolices 
are shown in Figs. 5 and 6. (in this connection note evidence 
given below under the excretory system that the latter are incom- 
plete). Although the foregoing facts point to possibly occasional 
augmentation in the number of proglottides in this region in adult 
worms, the usual appearances are as described below." And further 
it was noted that not only do the auricular appendages of the 
posterior ends of the proglottides disappear at about the same 
place, namely at about the 23rd or 24th segment, but that "there is 



80 

a more or less definite point in the strobila, at or about the 
15th proglottis, ahead of which the genital organs do not seem to 
develop and behind which in older strobilas they appear very 
quickly." Both of these facts pointed in the opinion of the 
writer to a more or less definite and predetermined number of seg- 
ments. Now the matter is cleared up considerably by the discovery 
that segmentation in this species is carried on after an entirely 
novel plan, involving the formation not only of new segments but 
whole chains of them or, in fact, strobilas from the original 
larval or primary strobila, as the writer will call it. 

The original larva (Fig. 10), quite comparable to the 
bothriocephalid plerocercoid, excepting for the perculiar scolex, 
gradually elongates with growth, until between a length of 4 and 
5mm. the first traces of segmentation appear in the hinder ends of 
cleared specimens as feeble aggregations of nuclei forming faint 
dark lines at regular intervals. In one specimen 4.8mm, in 
length five of these could be made out by close scrutiny, the 
second last of whichV/as 0.37mm. in length by 0.30 in diameter, the 
last one being slightly larger and rounded posteriorly. These 
primary segments elongate with the growth of the strobila while 
the constrictions between them gradually deepen as their anterior 
and posterior ends enlarge slightly, the former relatively faster 
than the latter. Fhen a total length of strobila of about 10mm. 
is reached the hindermost segment, itself now about 1.5mm. in 
length, begins to show faint transverse lines in its anterior end, 
decreasing in intensity from ahead backwards. These are the 
earliest traces of the divisions of the primary segments into the 



31 

secondary segments or definitive joints of the anterior ends of the 
adult strobilas. In other words the original or primary larva, 
plerocercoid or strobila divides up into secondary strobilas which 
eventually separate from each other and grow into the adult chains 
as described for the species. But long before .separation takes 
place the entire development of the anterior segments with their 
characteristic posterior auricular appendages and the formation in 
particular of the first segment can be followed with a consilerable 
degree of satisfaction in these primary strobila3 (Fig. 12). 
Whereas the writer originally (1914b : 32, figs. 5 and 6, PI. V) 
dre'f/ attention to young scolices with only 5 to 8 segments, he 
found in connection with the present study that the latter number, 
about 8 in external view or 16 or 17 in cleared specimens, is that 
developed by the secondary 3trobilas before detachment from the 
original chain. The smaller strobilas are now looked upon as having 
been prematurely and accidentally separated from the posterior end 
of the original or primary strobila. The attachment 30on becomes 
very slight owing to the rapid deepening of the construction ahead 
of the first segment, and some time before the auricular appendages 
of the latter are fully delimited posteriorly, very little manipula- 
tion of even alcoholic specimens, let alone cleaned one3, suffices 
to break up the chain. The writer, however, succeeded in finding in 
the material at hand one primary strobila, 88mm. in length, showing 
twenty secondary strobilas including the undifferentiated anterior 
segments from which they are developed. Furthremore, the last two 
of these, 10.4 and 11.5mm. in length, showed in their posterior un- 
segmented portions the earliest traces of the rudiments of the re- 
productive organs. As has been already 



82 

intimated, the anterior segments form within the secondary or 
definitive strobilas by a gradual demarcation from ahead backwards, 
first internally in the parenchyma, actually as transverse layers 
of nuclei (Fig. 12) which will eventually form the posterior auri- 
culate appendages, and then externally as sLown in the figure. 

A continued search for evidence in connection with the 
question of whether or not there is a definite number of segments, 
external and genital combined, brought out further interesting 
fact3. The number was counted in several young strobilas evi- 
dently not long separated from the primary 3trobila with the 
following results in the case of four typical specimens: (l) 
Length, 19mm., number of segments, 45; (2) 27mm., 29 or 30; (3) 
26.5mm., 30 (the posterior ones here ripe as in the next specimen); 
(4) 41mm., 32 segments. It would seem from these data that there 
is a more or less definite number of segments, which might be con- 
sidered to be about 30. But in No. 3, segments 9, 10, and 11 
were much elongated and show the earliest traces of further sub- 
division; while in No. 4 segments 10, 11 and 12 are likewise elong- 
ated and show not only similar traces of subdivision but particu- 
larly in the 11th early stages in the 3ame formation of secondary 
strobilas as described above for the primary strobila in its 
posterior end! Similar elongated segments in other strobilas 
show this condition near their middles instead of posteriorly; so 
that here we have a tertiary subdivision which must be considered 
as by no means as regular as the secondary subdivision of the 
original primary segments. These facts explain, then, the aberrant 
nature of the strobila in this region, noted formerly by the writer 



33 

(vide supra), and the presence in material of chains showing 
anteriorly very young segments similar to those formed in the 
oldest attached secondary strobilas but posteriorly much older 
segments with well developed auricles and farther back the typical 
mature proglottides of the ordinary strobila. Consequently we 
must consider that there is not a definite number of segments 
formed but that further, irregular and evidently indefinite sub- 
division, resulting in the formation of an inconstant number, 
takes place chiefly in the middle portion of the anterior segmen- 
ted region of what now must be called the secondary strobila. 

As to where all of this development takes place the 
writer has not come to any definite conclusions, knowing as yet 
practically nothing of the life-history of the species, for, 
altho all stages may be found in the intestines of Ami a calva , 
some of it may take place in the intermediate host. In a few cas- 
es primary strobilas showing three or more well developed secondary 
strobilas were found tightly coiled and surrounded by material 
which may have been from the intermediate host, whatever that is. 
At any rate it was of too firm a consistency to be merely coagulated 
mucCus from the host's intestine. 

On the other hand, this method of segmentation will now 
explain 3ome facts in connection with the nervous and excretory 
systems that were previously considered very unique j to say the 
least . 

In primary strobilas, even the youngest (Fig. 10) the 
excretory system consists, as in the adult, of a larger median 
vessel and two lateral vessels which run backward and unite in 



84 

the posterior end to form a plexus from which very many small 
vessels pass to the exterior by prominent foramina secondaria 
piercing the cuticula, much as described by Fraipont (1881 : 11, 
Fig. 7, PI. II) for Both r iocephalua scorpii . In the younge3t 
larva, that shown in Fig. 10, only the median vessel, which be- 
comes greatly reduced in diameter about twice the length of the 
bulbs from the anterior end, is present in the scolex. It forms a 
simple plexus among the bulbs anteriorly. In primary strobilas, 
however, in which segmentation has gotten well under way, all three 
vessels are quite prominent even in good toto preparations, and 
pass close to each other as well as to the chief nerve strands, 
through the constrictions between the developing secondary strobi- 
las, where the median vessel i3 somewhat enlarged. As they near the 
anterior end of the worm they give off numerous branches of their 
own calibre, and when they meet the large ganglionic masses, 
described below, diverge as four vessels (two on each side) and 
continue lateral to the bulbs to the tip of the scolex. Here, 
after forming an open plexus among the anterior ends of the former, 
they unite in a single median frontal loop. As the constrictions 
between the secondary strobilas deepen all three vessels likewise 
become gradually constricted until eventually they are cut off, 
and the adult conditions are subsequently developed by a process 
of turning in of both severed ends. And since the median vessel 
was considerably enlarged at the region of constriction, it remains 
thus in the hinder end of the adult strobila, as well as in the 
first segment (former 3colex) as described and figured elsewhere 
by the writer (1914b : 93-95, Figs. 13, 37), while in the latter 



85 

it is joined by the lateral vessels to form the characteristic 
terminal vesicle. , 

The nerv~ous system of the primary 3trobila consists of the 
two chief nerve strands passing thruout the segments, a quite 
irregular commissure connecting them anteriorly, and a very large 
ganglionic mass, situated some distance posterior to the proboscis 
bulbs. The chief nerve 3trands, which the quite indistinct :--t 
different levels but constantly located in the median frontal 
plane, diverge as they meet the ganglionic mass in passing forward, 
and consequently opposite the bulbs come to lie close to the sub- 
cuticula laterally (Fig. 15). About 0.3mm, from the tip of the 
scolex they are united by a very irregular but comparatively large 
transverse commi 33uce,irom which large trunks pass to the neighbor- 
ing bulbs both forward and backward. This commissure \vas found 
to be present wholly or in part in about 12 sections of an 8jul 
transverse series, hence its length or longitudinal diameter is 
about 0.10mm., while its maximum depth between the latsral pairs of 
proboscides is about 40 ^ . Large branches are given off anteriad 
to the lateral walls of the bulb3 and caudad to the central walls. 
In the latter case a large branch was found to leave the median 
portion of the commissure, which i3 incidentally freely pierced 
with excretory vessels, on each surface and shortly divide", into 
two, each supplying the central walls of one of the frontal pairs 
of bulbs dorsally or ventrally (Fig. 15). The anterior branches 
likewise arise in a common trunk on each side, which is in reality 
the continuation of the lateral ganglionic enlargements of the 
commissure, but they supply the outside wall of the lateral pairs 



86 



of bulba. Imbedded in the commissure were seen numerous nuclei 
which, on account of their larger 3ize than the neighboring paren- 
chymatous nuclei, were interpreted as ganglionic or nervous in 
their nature. Numerous among the posterior end3 of the bulb3 and 
extending far from 0.8 to 0.9mm. backward beyond them (Fig. 11) 
fee re i3 present a large mass of large nucleated cells which in 
transverse sections (Fig. 16) are seen to occupy the whole of the 
meiujlla^and about the whole of the section) excepting for the ex- 
cretory vessels. These cells are roughly spherical in shape and 
have a maximum diameter of 25^., their nuclei being 5^. On 
account of their finely granular consistency and their taking the 
orange- counter 3tain quite as does the anterior nerve commissure 
they were interpreted a3 being ganglionic cell3. And this view 
•vas supported on closer study by the discovery that they are not 
only united laterally with the chief nerve 3trands (Fig. 16), 
which can scarcely be distinguished from them at various levels, 
but with each other thru a complicated plexus of fine longitudinal 
strands which pass forward towards the bulbs and form around their 
bases an almost solid mas3 of fibrils (Fig. 11). From this mass 
large strands from 10 to 15^ in diameter pas3 into the bases of 
the bulbs, one for each, and are distributed among the retractor 
muscles of the proboscis which they evidently supply. In the 
youngest primary strobilas, but not in the older one3, this mass 
of fibrils at the bases of the bulbs evidently connects forwards 
by a few strands with the commissure. 

Just as the definitive form of the anterior and posterior 
end3 of the excretory system is explained by the separation of the 



37 

secondary strobilas and the subsequent healing over of the cut 
ends, so i3 that of the nervous 3ystem, particularly anteriorly. 
In the original description the writer (1914b : 93) said that: 
"The nerv^ous system consists of a nerve-ring situated immediately 
beneath the tip of the scolex and covering the median excretory 
vesicle like a cap, and the two chief nerve strands passing back 
from it through the whole of the strobila. The former ia a com- 
paratively weakly developed structure (Fig. 11), elliptical in 
transverse section, with diameters of 60 and 40p- As was shown 

in Fig. 11, the nerve-ring is drawn out forward into a point which 
is directly opposite a small conical pit in the tip of the "3colex", 
facts which formerly seemed rather strange to the writer in com- 
parison with conditions in other bothriocephalids . Now they, as 
well as the relatively small size of the nerve-ring, are explained 
by the contraction of the free end of the n 3Colex" after separation 
and the growing together of the ends of the nerve strands with 
subsequent differentiation into the nerve-ring. The close associa- 
tion of the nerve-ring and the terminal excretory vesicle is also 
quite comprehensible in the light of this method of development, 
for, 3ince the nerve strands are situated close outside the lateral 
excretory vessels at the constrictions, they simply turn in towards 
the median line and unite immediately ahead of the junction of the 
latter with the median vessel. 

As will be gathered from the foregoing description there 
is a most remarkable resemblance between the scolex of K. gl obuli- 
f orme and that of the Trypanorhyncha not only in the structure of 
the probo3cides but also in the presence of the large mass of 



88 



ganglion cells associated with them posteriorly. Each proboscis 
consists of three parts: (1) a hollow tentacle, capable of s vagi na- 
tion, (2) a short, permanently protruded stump, armed 7/ith thickly 
set minute cuticular spines, and (3) a comparatively elongated bulb; 
of which (1) and (2) may be comp-red respectively with the proboscis 
and the bulb of Tetra rhychus or Rhynchobothrius . The proboscis, 
altho not provided with any kind of armature, is nevertheless 
supplied with a group of well developed retractor muscles which is 
evidently analogous in function to the single retractor muscle of 
the Trypanorhyncha. The bulb is not only provided with a muscula- 
ture arranged so as to dmini3h on contraction the volume of the 
organ but is al30 lined with an epithelium like layer comparable to 
that of the members of the latter group. But 3ince the bulb extends 
to the point of exit of the proboscis, there is no part strictly 
analogous to the proboscis — sheath of Tetrarhynohus , altho the 
stump would at first sight seem to be such. Furthermore, the cells 
forming the large mas3 behind the bulbs in Haplobothrium , which are 
here interpreted a3 ganglionic cel^s, bear not a little resemblance 
to those described by Braun (1896 : 1294) after Pintner (1880), 
Lang (1881) and Niemieo (1888.,) as associated with the bulb3 of 
Tetrarhynchus longi colli s (v. Ben) (= D i b o t h r i o r hyfc hu s rufi colli s 
Monti celli ) and considered by 3ome to be ganglion cells and by 
others myoblasts. The distribution of the large nerve trunks 
arising from the nerve commissure is also somewhat suggestive of 
conditions in a few of the tetrarhynchids (cf . Braun, 1896 : 1293- 
94) . 

While the writer is not prepared to go further into this 



39 

comparison he would like to emphasize the significance of the 
layers composing the walls of the bulbs in H. glob uli forme in 
connection with the possible origin, of these most aberrant struc- 
tures. In discussing the homologies of the proboscides of the 
Trypanorhyncha Benham in Lankester's "Treatise on Zoology" (p. 127) 
said that, "It appears more probable (Pintner) that each proboscis 
has been developed by the deepening and modification of an 'acces- 
sory 3ucker» of some Tetraphyllidean, as its relation to the 
bothridia, and its mode of development, closely agree with these 
structures. Functionally, too, it is a perfection of the armature 
plus the accessory sucker of three forms [Aeanthocephala, lu'emertini 
and Taenioids}; whilst there is no doubt that the 'phyilidea* of 
the two orders are identical." The fact that here the walls of 
the bulb, since they are composed of an outer layer of longitudinal 
muscles, a middle layer of circular fibres and an inner cuticular 
layer, are not only quite comparable but directly continuous with 
the cuticula and cuticular muscles of the body wall and in the 
reverse order, would seem to lend sup;ort to Pintner' s view. For 
simple invagination of the external layers of the body wall in 
development would account for these structures, while the proboscis 
with it3 retractor muscles might well be formed by the modification 
of the external layers of an "accessory sucker". The writer has, 
however, no fact3 to support this theory since in the earliest 
primary 3trobilas or plerocercoids met with the prob03cides were 
already well developed. 



so 

Subfamily 3. C7ATH0 CEPHALINAE Luehe, 1899, e.p. 

Scolex unarmed, not longer than broad, with two surficial 
sucking grooves, more or less fused with onqjanother or one terminal 
one of a sucker-like structure. External segmentation little ex- 
pressed or quite absent. Genital organs in each segment simple. 
Genital openings surficial median. Vagina and uterus open into a 
common vestibule, — in young proglottides near on^fanother, — 
lying behind the male opening and similar to the genital atrium of 
other cestode3, which may be surrounded by a sphincter like muscu- 
lature. The genital openings of the differnnt segments do not open 
on the same surface of the worm, but alternate irregularly, being 
sometimes on the one surface and sometimes on the other. Uterus 
a coiled canal without uterus-3ac. 

Sexually mature in the intestines of fishes. 

Type genus: Cya thoceph alus Xessler 

The above is Luehe f s (1910 : 22) diagnosis modified to 
read "may be surrounded, etc." instead of "i3 surrounded, etc." 
in connection with the genital sphincter, since in 1he 3pecies 
described below no such structure was found. 

Genus 1. Cyath ocepha lus Ke33ler, 1358. 



Taenia (part) 


Pallas 


1781 


40. 


Taenia (part . ) 


Bot 3ch 


1786 


71,107 


Eehinorhychus (part.) 


Zeder 


1803 


291. 


Cephalocotyleum 


Biesing 


1850 


617. 


Qathocephalu3 


Kessler 


1833 


135 



qjathooephalua 
Honahothrium 
Aorobothrium 
Cyathocephalus 



Sri mm 

Tl 

Olsaon 

Z3chokke 

Loennberg 

Kraemer 

Olsson 

Luehe 

Braun 

Luehe 

Luehe 



1371 

1372 
1884 
1889 
1892 
1893 
1889 
1900a 
1900 
1910 



502. 

504. 

40 

37 

42 

548 

14 

53 

1697 

12 

22 



Generic diagnosis: Soolex a single, undivided terminal 
3ucking organ, which in its form and structure no longer shows an 
origin from two fused suriicial bothria. External segmentation 
only slightly indicated. Sphincter surrounding the female genital 
cloaca apparently little developed. 

Occurrence: In Teleo3t3. 

Type species: Oyathocephalus truncat us (Pallas, 1781) 



Cyathocephalus amsri canus sp.no v. 
? 1897 Cyathocepha lus truncatus Linton, 1397 : 438-29. 

Specific diagnosis: With the characters of the genus. 
Small cestode3, up to a length of at least 11mm. with a maximum 
breadth of 1.1mm. Scolex funnel-shaped, 0.3-0. 5mm. long and 0.5- 
0.7 broad, with re volute edges. Neck 1.0-1. 8mm. in length. Seg- 
ments twice a3 broad a3 long, terminal one rounded. 



92 



Cuticula 5juu in thickness, with no hooka nor spines; 
3ubcuticula 35 to 40^ . 

10 to 13 sets of genitalia, beginning 1.5 to 2.0mm. from 
the anterior end. Strong tendency for the reproductive apertures 
to lie all on one surface of the 3trobila. Vagina opens behind 
uterus. Neither papillae nor sphincters around the genital 
openings . 

Testes in two laterial fields in the medulla of the an- 
terior portion of the proglottis, 60 in diameter. Coiled vas 
deferens anterodorsal to cirrus-3ac; no seminal vesicle before 
entering cirrus-sac nor connective tissue sack surrounding the 
whole duct. Protruded cirrus 0.2mm. in length by 0.13 in diameter 
at base . Cirru3-sac ovoid in shape 0.20-0. 33mm. in length by 
0.17 in diameter; no retractors connecting it with the dorsal body 
wall; large ma33 of glandular pigmented cells surrounding it 
dorsally and laterally. 

Vagina 15 jul in diameter; no sheath near it3 opening; re- 
ceptaculum semini3 75^ . Spermiduct very short and narrow, 35 and 
8 ja- respectively. Ovary tubulo lobular, fan-shaped; wings extending 
dorsally and laterally around the ventral genital duct 3; isthmus 
prominent, 0.18 x 0.10mm.; ova in same 13-15 in diameter. Oo- 
capt 25^ in diameter. Vitelline follicles continuous from pro- 
glottis to proglottis, forming a layer 70^ thick in the cortical 
parenchyma, 30 to 35 y. in transections. She 11- gland dorsal, 
voluminous. No muscular 3ack surrounding the uterine rosette. 

Eggs , 40 x 30 J*/ . 

Habitat: In stomach, pyloric coeca and intestine of the 

host . 



33 



Host Locali ty Collector Autho rity 

Coreg onus olup eiforml3 Outer Id., J. W. Milner Linton, 1897TD : 

L.Superior 429. 

" " Off Giant's Cooper Cooper, (the 

Tomb Id., present paper). 

Georgian Bay 

Type specimen : No. 165A, C. A. R. C. 

Co- type : Ho. 1853, C. U. 111. 

Type locality : Georgian Bay, Lake Huron, off Giant's 

Tomb Island. 

Altho the species described here is closely related to 
0. truncatus of Europe and evidently the same form was reported 
as such by Linton (1897 : 428) from the 3ame host in which it was 
found by the writer, it preaent3 30 many differences, even barring 
some probable errors by Kraemer (1892), from that 3pecie3 that it 
ka here considered to be new. 

A3 shown in the appended table where the largest specimens 
at hand are dealt with, this 3pecies is considerably smaller than 
the European species which ranges from S to 40mm. in length by 
1.5-4 in width. Linton gave these measurements as 7 and 1.3mm. 

The general shape of the body, however, is the same as are 
the proportions of the 3C0lex and proglottides. The border of the 
infundilobif orm scolex (Fig. 18) is thickened and almost constantly 
rolled backward 3omewhat as in the figures given by Zschokke 
(1834a, Fig. 9] and Kraemer (1893, Fig. 5), the funnel being about 
0.33mm. in depth and usually filled with a plug of mucous membrane 
from the V.ost's alimentary tract. The posterior limits of the 
scolex are difficult to define since the organ gradually narrows 



94 

down and then as gradually enlarges again to form the neck. The 
latter was considered to include that portion of the anterior end 
of the worm between the narrow--- 3t region behind the 3Colex and the 
first vitelline follicles which are situated 3ome distance ahead 
of the first cirrus-3ac. The maximum breadth of the 3trobila la 
at the posterior end of either the first third or one half. The 
segments are, as described by various writers for 0. truncatua, 
about twice a3 broad as long, the last one, however, being rounded 
posteriorly and provided with a notch in the middle which accommo- 
dates the exi^t of the excretory vesicle. They are, furthermore, 
closely united, as pointed out by Linton (1897 : 429) when he 
said that, "The bodies of these specimens appear to be unsegmented, 
or, at least, with only very faint indication of division into 
segments." A3 a matter of fact numerous transverse wrinkles pre- 
sent in mo3t specimens make it almost impossible without the 
external evidences of the inner genitalia to|iistingui3h the limits 
of the proglottides. And in thia respect they agree with C. 
truno atua, 3ince Zschokke (1084 : 38) said concerning the segments: 
"11$. aont 3olidement fixes les una aux autre3, leurs limites aont 
iiff icilement vi3ibles." The following table gives the measure- 
ments of four of the largest specimens'. 



95 



Number 


165.4 


165.5 


43.1 


165.1 


Length 


10mm . 


9mm . 


11 mm . 


7 . 5mm . 


Maximum breadth 


1.01 


0.92 


1.11 


1.05 


Length of neck 


1.48 


1.00 


1.48 


1.8 


Breadth scolex, tip 


0.55 


0.53 


0.74 


0.64 


> base 


0.37 


0.42 


.55 


0.30 


Length of scolex 


0.48 


0.33 


0.61 


0.50 


Number of sets genitalia 


13 


12 


13 


10 


First cirrus from ant. end 


1.85 


1.56 


2.25 


2.01 


Remarks 


Toto 


Toto 


Toto 


Sectioned 



The cuticula ia 5jt in thickness over the 3Colex as well 
as on the segments and divided into tvfo layers, the outer of which 
is about one-half as thick as the inner and mors or less irregular 
in structure. In most places, in fact, it was found partially 
or wholly separated from the inner and more homogeneous layer and 
in such a manner a3 to present a distinctly wavy appearance. 
However, no 3uch chitinous hooks as described by Kraemer (1892 : 
10) for thejcuticula of the lateral borders were seen, but the whole 
tissue is freely pierced with numerous foramina secundaria of the 
excretory system which in C. tru nca tu3 Eraemer considered to be 
the points of entrance of nutriment. The thickness of the cuti- 
cula, according to the same author, is ISy* , an outer irregular 
layer being 5|^and showing a sort of ecdysis ( "Haatungst roce3s") . 
This, however, may be simply the separation of the outer layer of 
the cuticula from the inner as mentioned here, since h^s^aid "Die3e 
Auffassung wird dadurch erhartet, dass sich an einigen Stellen 



96 

die3er Belag nicht fin - ;et, dafur eine yunge homogene Cuticula," 
the latter being then the inner homogeneous layer. At any rate 
it is quite evident that the cuticula of 0. t rune a t ug is a much 
thicker tissue than that of the form described here, — and no 
one else than Kraemer seems to have described it. 

The subcuticula varies in thickness from 25 to 40(^ , being 
thus quite comparable to that of C. trun catus in which it is 38 ^ . 
The elongated, quite columnar cells of the same were found to be 

in diameter, a3 were their comparatively large nuclei, the 
respective measurements by Kraemer being 19 and 6p. . 

On account of the fact that the material studied was not 
as well fixed as the writer would like to have had it, the paren- 
chyma and likewise the calcareous bodies were quite unsatisfactory 
from an anatomical standpoint. The spaces which were considered 
to have accommodated the latter, before they were dissolved out 
during fixation, — and in some of them the nuclei of the bodies 
were still visible, — were in general ellipsoidal to almost a 
spherical in shape and from 13 to 25 pi in length by 7-18 in width. 
According to Linton they are 10-20 ^ long, while Zschokke gave 
them as from 8-10 p. , and Kraemer, 30 x 18 ja. . 

In general the musculature is as described by Kraemer, but 
all the groups are comparatively weakly developed (Fig. 19), the 
longitudinal layer, for instance, being only 20 pu in thickness in 
the median line posteriorly and about 60 ^ in the neck region 
(76j^ in C. trunc atus ) where the dorsoventral and transverse fibres 
are also much stronger than elsewhere. In the anterior part of 
the neck, particularly immediately behind the 3colex, the fibres 



3? 



of the two latter series are much stronger, altho less numerous 
than farther back. A3 they pass the posterior end of the funnel 
they become circularly disposed, longitudinally as well as trans- 
versely, and from there on to the tip of the scolex gradually more 
numerous close around the funnel of the organ, for the control of 
which they obviously act as constrictors. Antagonizing these are 
numerous weaker radial fibres, arranged as in C. catena t us 
Riggenbach (= Diploootyle Rudolphii Mont. — Luehe 19C3a : 330) 
where they were considered (Riggenbach, 1898 : 639) to be derived 
from the longitudinal muscles with which they are continuous at 
the base of the scolex. Altho they mingle freely, yet quite 
separately, among the latter, the writer is inclined to the same 
view regarding their homologies in C. americanua , since it is quite 
evident that the dorsoventral and transverse fibres, which might 
otherwise be considered to give rise to them become modified to 
form the circular muscles of the scolex. And this view if further 
supported by the fact that only a very few of the longitudinal 
muscle e of the neck pass for a short distance beyond the bottom 
of the funnel, but most of them are inserted in the latter, thus 
functioning with the radial fibres in enlarging the organ of adhe- 
sion. As pointed out by Riggenbach these radial muscles are 
apparently absent from C. truncatus ; they were not described by 
Kraemer,but the enlargement of the funnel was considered to be 
accomplished by the contraction of the dorsoventral fibres (cf . his 
Fig. 1). The outermost layer of circular and longitudinal muscles 
in the scolex, which are merely extensions of the cuticular muscles 
of the neck region, are not nearly so strongly developed here as in 



38 



£. truncatus, so far as the writer can gather from a comparison of 
the material studied with Kraemer's description. There is, however, 
in the neck region, particularly in its anterior portion, a series 
of outer longitudinal muscles which, while situated in transections 
among the outer clear ends of the subcuticular ceills and very close 
to the longitudinal cuticular fibres, are nevertheless quite dis- 
tinct from the latter. At the base of the 3colex they pass in- 
wardly between the cells of the subcuticula and continue farther 
towards the anterior border of the funnel than do the inner longi- 
tudinal fibres. Posteriorly they diminish considerably in number 
but may readily be seen in the mature proglottides. 

The nerv.ous system is arranged in general as in C. trunca- 
tus, but the longitudinal trunks are only 36 thick by 13 wide 
(0. 345mm. according to Eraemer). They are 3ca*cely enlarged 
anteriorly to form ganglia, as shown in Kraemer*s Fig. 5, but are 
each divided into two distinct dorsoventral halves which gradually 
diverge as they pass on into the walls of the funnel to form four 
large nerves. No single transverse commissure connecting the main 
trunks behind the cavity of the 3colex was seen but rather a number 
of fine cross-connections which were not made out satisfactorily. 

On account of the poor preservation of the parenchymatous 
ground tissue of the material studied the excretory system was not 
brought out in actions to the writer' 3 satisfaction. All that can 
be said in this connection is that there is an inconstant number of 
longitudinal vessels in transection, evidently more than the six 
of _C. truncat us , which do^riot occupy definite positions but anasto- 
mose freely with each other especially in the lateral portions of 



the medulla. In the 3colex these vessels are smaller and the 
anastomoses are much more numerous, while posteriorly at least 
two pass into a quite irregularly shaped terminal vesicle, which 
however in the light of Wolf's (1906) finds cannot be considered 
to be a true terminal excretory vesicle. As above stated, 
foramina secundaria are quite numerous in the cuticula. 

The reproductive organs appear quite close behind the neck, 
the vitelline follicles being situated from 1.3 to 2.3mm. from the 
anterior border of the scolex, and the first cirrus-sacs from 1.6 
to 3. 2mm. From 10 to 13 sets of genitalia were observed for this 
species. These follow each other closely and are not separated by 
any 3epta or other boundaries, the vitelline follicles being, in 
fact, strictly continuou.3 from proglottis to proglottis. The open- 
ings of the cirrus-sacs vary from 0.45 to 0.75mm. apart, but, as 
pointed out by Kraemer, these measurements are of little diagnostic 
value on account of the different states of contraction. As in 
C. truncatU3 the reproductive openings are all on one surface of 
the proglottis but alternate as a unit irregularly from one surface 
to the other, there being, however, a strong tendency for them tc 
lie all on the one face of the strobila. This alternation also 
involves the ovary, the isthmus of which is arbitrarily considered 
to be ventral. It usually lies on the same surface as the repro- 
ductive openings, so that when the latter passes to the opposite 
surface it moves accordingly. This alternation of the openings 
has, of course, been known ever since Pallas described Taenia 
t run cat a in 1781, but, so far as the writer is aware, no one has 
dealt with the relations between the openings and the ovary noted 



. 100 

here. Concerning thi3 matter Kraemer said only: "Das Vernal ten, 
da3S die Geschlecht3organe alternirend dorsal-und ventral nach 
aus3en mimden, erinnert in gewisser Beziehung an die alternierende 
marginalen Ge3chlecht sof f nungen verschiedener Fisch- und Vogel 
taenien, und wurde bereit3 von der er3ten Beobachten, Pallas und 
Bat sen erkannt, d.h., 3ie hatten auf ceiden Flachen die fortlaufen- 
de Peihe der 'Punkte' wahrgenommen, ohne sie indessen als Aus- 
mundungen der Sexualorgane zu deuten. Die neueren Beobachter haben 
sammtlich dieses oben be3chriebene Verhalten ufcersehen, und geben 
die Geschlechtsof fnungen als ventral gelegen an." But it should 
be mentioned here that whatever is the stimulus which, during the 
very early stages of developmant of the sets of genitalia from 
their nuclear rudiments, causes the reversal of the "'hole proglottis 
it would seem to be such at times as to fail to bring about the 
turning over of all parts of the rudiment. As shown in Fig. 20, 
which is a diagram of a sagittal series of seven proglottides in- 
cluding the endlone, the cirrus and female genital cloaca of No. 3' 
from the top have gone to the opposite surface while the ovarian 
isthmus, represented by the solid black disc in each segment, ha3 
remained on the same surface as those in segments 1, 2 and 4 in 
the immediate neighborhood. Here the stimulus which brought about 
the uni surf ici ali ty of the latter may have influenced the ovarian 
portion of the common rudiment of No. 3 and caused it to lag behind, 
while the more peripheral rudiments of the cirrus, vagina and distal 
portions of the uterus were freer to move. This arrangement of 
the parts in the aberrant segment in question naturally caused con- 
siderable departures in the courses of the reproductive ducts from 



101 

the normal, as were observed. 

The genital openings were found to vary from 75 to 115 (A 
apart, but, as pointed out by Kraemer, these data are of very 
little specific value. The vagina and uterus open in a common 
genital senus or cloaca, but unlike conditions in the European 
species the vagina opens constantly behind the uterus and slightly 
to one side and not ahead of it. Furthermore, neither papillae 
nor sphincter muscles were found around either or both genital 
openings in this species. The female genital cloaca, usually 
situated at the bottom of a depression and often on a low papilla, 
ranges from 30 to 60 p in depth. In frontal sections it is seen 
to be in the form of a transverse slit about 60u, in length, into 
the ends of which the vagina and uterus empty; that is, the vagina 
opens diagonally behind the uterus and usually to the right of it. 
It is lined by a direct continuation of the cuticula from the sur- 
face of the segment. The general habit of the reproductive organs 
is shown in Fig. 21 of a frontal section of a mature proglottis. 

The testes are situated in the medu^llary parenchyma in 
two fields lateral to the cirrus-sac, or more strictly speaking, 
in the lateral portions of the region between the cirrus-sac and 
the ovary of the proglottis ahead, since they extend forv/ard to the 
latter and backward to the anterior ends cf the wings of the ovary 
of the segment to which they belong. They are noteworthy on account 
of their clear appearance (Fig. 19), as in 0, truncatus , very little 
contents having been seen in all of the sections made. While their 
3hape is usually spherical or somewhat flattened enteroposteriorly 
according to the condition of contraction of the segment, their 



102 

maximum diameter was found to be about S0f/L . The coila of the 
vas deferens, altogether about 0.30mm. in diameter are accommodated 
in the somewhat confined space dorsal and anterior to the cirrus- 
sac, extending to the ovary ahead (Fig. 31). Whereas Kraemer gave 
the diameter of the duct as 0.133mm. or about eight times as much 
as just before it enters the cirrus-sac, it was found to be only 
44 y at the most in this species. Furthermore, it was not found 
to be enlarged to form a 3eminal vesicle close to the cirrus-sac, as 
shown in Kraemer' s Fig. 6 and 13, but to gradually diminish in 
size until as it pierces the wall of the latter it3 diameter is 
only 10 ^ . Nor is the whole vas deferens enclosed in a connective 
tissue sack, such as described by Kraemer. Within the pouch it 
enlarges considerably to form a thin-walled inner seminal vesicles 
which is quite conspicuous in sections, situated for the most part 
nearer the proximal end of the former but often lying alongside the 
cirrus proper. It may attain a diameter of 50 \l even when empty, 
— in none of the sections made was it found to contain spermatozoa, 
these having been probably extended at the moment of fixation. 
Then follows the cirrus proper which is sharply separated from the 
3eminal vesicle, since it protrudes backward into the latter with 
a diameter of 10fA. and for a distance of from 15 to 35 ^ • The 
cirri were found protruded in most of the specimens at hand, in 
which case they had a maximum length of 200 jul , diameter at the 
base, 120 |m , and at the tip about 40 f*- . The thick cuticula cover- 
ing the organ is decidedly roughened or irregularly "cleft", es- 
pecially towards the tip, but not provided with spines of any kind. 
Incidentally, sections show that the protrusion of the cirrus, on 



103 

account of its size, results in the eversion of almost the whole 
of the contents of the sac. The length of the cirrus within the 
sac is at least 185/a- , — it is usually bent once in its proximal 
portion, — while its diameter varies considerably. The layer of 
parenchymatous and myoblastic nuclei surrounding the cirrus within 
the sack is about IOjjl in thickness as compared with 5yc in C. 
truncat us . In sections of the extended cirrus most of these 
nuclei appear in the tip of the organ surrounding a good deal of 
the cuticula which still remains invaginated; but they are in all 
probability myoblastic as are others farther back along the course 
of the retractor fibres. In frontal sections the cirrus-sac is 
circular in outline (Fig. 21) with a maximum diameter of 175 f*. , 
while in transverse and longitudinal sections the depth ranges from 
300 to 230 Jul > the whole structure being somewhat ovoid in shape 
with its slightly smaller end directed outwardly. Its wall is 
comparatively thin, ill defined and composed of a somewhat loose 
network of muscular fibres running irregularly obliguely in 
all directions, 30 that sections cut in any plane show them almost 
circularly arranged. With emission of + he cirrus the wall is 
fairly difficult to locate, since its innermost fibres are not 
easily differentiated from the retractors of the cirrus proper 
which bulk, largely in the contents of the sac and since it is not 
provided with any dorsal retractors connecting it with the dorsal 
body wall as described by Kraemer for C. trunca tus . Forming a 
sort of gland closely applied to the wall of that part of the 
cirrus-sac within the medulila there is to be seen, even in toto 
preparations, a comparatively large mass of large darkly pigmented 



104 

polygonal cells (Fig. 21). In frontal sections they lie on each 
side of the sac, not extending- much beyond its anterior and pos- 
terior edges, the whole structure being thus shaped some what like 
a saddle. Each cell is elongate in shape, provided with a well 
defined wall, prominent the not especially large nucleus, and very 
granular and highly pigmented cytoplasm, the color of the pigment 
being dark brown. Altho they are very closely arranged around the 
wall of the cirrus-pouch and most of them are quite pointed towards 
the 3ame ; their function is pretty much a matter of conjecture, 
unless perhaps they are the much modified myoblasts of the muscles 
of the walls of the pouch, which is suggested by the relations of 
the inner attenuated ends of some of them with the latter. No 
such cells have been described for the European species, so far as 
the writer is aware; but here it must be emphasized that they are 
very conspicuous. 

From its opening which has been dealt with above the 
vagina proceeds dorsally almost at right angles to the surface of 
the proglottis, and then within the medulla turns backward with a 
few coils to expand into a comparatively enormous receptaculum 
seminis which on account of its size, can scarcely be distinguished 
from one of the coils of the uterus unless it is traced out. At 
the turn in it3 course it has a diameter of about 15 y~ and is lined 
with a continuation of the cuticula of the female genital cloaca, 
5 in thickness and surrounded by a layer of circular muscles. 
4a it passes above the ovarian isthmus its cuticular lining gradu- 
ally diminishes in thickness so that the seminal receptacle is 
provided with a very thin layer only. While the latter may have a 
diameter of 75a slightly behind the isthmus of the ovary, it 



105 

narrows down very abruptly to a very small 3permiduct, 8 jtt in dia- 
meter and about 35|a. in length before joining the oviduct. In 
distinct contract with C. truncatus there is no "connective tissue 
and muscular sac" surrounding the beginning of the vagina as de- 
scribed by Kraemer, but only the usual mass of nucleu most of which 
are subcuticular in their nature. The ovary (Figs. 19, 21) is a 
tubulolobular organ, the limbs of which extend fanwise laterally 
and dor sally from the ventral isthmus anteriorly as far as the 
cirru3-3ac and dorsally thruout the whole of the medulla, thus 
surrounding the central connections of the genital ducts and the 
coils of the uterus (Fig. 21). The wings in whose irregularly 
shaped tubules young ova in various stages of development are to be 
seen, connect with the rounded isthmus by narrow portions quite as 
described and figured by Kraemer, altho he evidently erroneously 
called the isthmus the "oo^typ . " The latter in 0. americanus 
has a width of 0.18mm. by a length of 0.10 as compared with the 
similar me surements of 0.19 and 0.07ram. in the cases of C. trun- 
Oatus . Ova from the isthmus measured from 13 to 15 //- in diameter, 
their nuclei 7-8 and their nucleioli 4, those of the latter species 
being 9-13 according to . Grimm (1871) and 15 (jl according to 
Kraemer who gave the diameter of their nuclei as 9^ . The oviduct 
begins with a rather 3hort oocapt (Fig. 22), 26 pi in diameter, and 
proceeds for only a comparatively short distance, with a diameter 
of from 15 to 20 , before being joined by the spermiduct. A 
little farther dorsally it i3 met by the vitelline duct which 
comes from the ventral portion of the medulla just ahead of the 
isthmus where it is formed oy the union of a right and left duct as 



in C. truncatus . Thruout its dorsoventral course it is expanded 
with a vitelline reservoir which may reach a diameter of 40^o , 
especially ventrally. Immediately outside of the longitudinal 
muscles the vitelline follicles form a compact layer about 70 
in thickness (l.~3p- in C. tru ncatus ), continuous from proglottis to 
proglottis and broken only immediately around the reproductive 
openings. In frontal sections where they are cut tranversely their 
greatest diameters being at right angles to the surface of the pro- 
glottis, they range from 30 to 85 . since they are usually some- 
what flattened anteroposteriorly . Their number in transverse sec- 
tions baries from 2£ to 35, 45 being given by Kraemer. From it3 
point of origin to a short distance beyond the entrance of the 
vitelline duct, the ovident is lined with epithelial cells with 
prominent nuclei but indistinct boundaries, so that the whole is 
of the nature of a synoitium. But soon this epithelium becomes 
modified so that, as the duct continues with a few coils to the 
opposite side of the proglottis, its walls are quite thin yet 
clearly nucleate. Then as it further enlarges dorsally it becomes 
surrounded with the voluminous 3hell-glan& which, follows the tube 
thru a number of its coils (Fig3. 19 and 31) even on to the begin- 
ning of the uteru3, for in frontal sections about one half of the 
posterior half of the uterine rosette i3 surrounded by them (Fig. 
21). The cells compo3ing the gland are comparatively short, stout 
and well defined, their nuclei large with the nucleoplasm quite 
clear like the cytoplasm. No shell-gland such as described by 
Kraemer was found in this species. Beyond this elongated ootype 
the duct continues as the uterus proper which has its largest coils, 



107 

up to a diameter of 0.10mm. or more when filled with eggs (0.033mm. 
in C. trun catus) , just behind the cirrus-sac. Before reaching the 
opening, the position of which has been stated above, the tube 
narrows down quickly. Thruout its course it is lined with a much 
attenuated epithelium, the nuclei of which, however, stand out 
prominently towards the lumen. In this species there is no mus- 
cular sack surrounding the uterus, as described and figured by 
Kraemer . 

The largest eggs in the uterus not in a collapsed 3tate, 
were found to be ellipsoidal in shape and 40 x 30 ^ in size. Linton 
gave the 3ize when preserved in acetic acid as 50 x 32^ ; while the 
measurements for C . truncatus have been given as S5 x 76 (a (Kraemer) 
and 44-51 x 33-36 [a. (Luehe, 1910). Since most of the eggs seen in 
the uteri of the setions made were quite young, many of them not 
having gone thru the first cleavage as yet, the writer is of the 
opinion that the size of the egg of this species is probably the 
same aa that given by Luehe for C. trun catus in Europe. 

Altho, so far as the writer is aware no one has as yet 
studied the early stages in the development of C. t i/uncatus , Wolf 
(1906) discovered that the intermediate host i3 Gramma ru 3 pal ex 
and that the transfer to the final hosts is a direct one. As re- 
gards C. america nus the writer is unable to name the intermediate 
host, but he is of the opinion that further work will in all pro- 
bability show that in Pontoporeia hoyi (Stimpson Mss.), at least 
in Georgian Bay where it is found in large numbers as practically 
the only food of Cor e g onus olupeiformis in which the parasite was 
found (vide Huntsman, 1915 : 150). 



108 



In the above description it has been shown that this species 
differs from the .veil known C. trunca tus of Europe in a great many 
points, but in none so radically as the following: The absence 
of chitinous hooks on the cuticula.. of the lateral borders; the 
presence of radial muscles in the walls of the 3colex and of a 
number of fine nerve commissures connecting the chief nerve strands 
anteriorly instead of a single one; the vagina opening behind the 
uterus opening; the absence of papillae and sphincter muscles sur- 
rounding the genital openings; no enlargement of the vas deferens, 
to form a seminal vesicle ju3t before entering the cirrus-pouch; 
no connective tissue sac surrounding the <vhole of the coiled vas 
deferens; the absence of dorsal retractor muscles of the cirru3-sac, 
and the presence of the peculiar glands closely surrounding the 
same; no "connective tissue and muscular sac" surrounding the be- 
ginning of the vagina; the very different central connections of the 
genital ducts as regards the ovarian isthmus ("ootyp" of TCraemer); 
and, lastly, the absence of any such "shell-gland" as described by 
the 3ame author. Consequently the writer considers this the 
American form to be specifically different from the European form 
and proposes the name Cyath oc ephal u3 americanus sp.nov. 

The material studied consisted of two lots, IT03. 43 and 165 
of the writer's collection from the stomachs of several specimens 
of Co regonu s clupeif ornii s (Mitchell) from GeorgianBay, Lake Huron, 
as listed above . 



109 



Subfamily 4, MAR 3 1 P ME T R I N AE subfam.nov. 

Scolex with two typical and fairly deep bothria and a 
[ terminal disc. External segmentation very distinct and regular. 
Opening of cirrus and vagina marginal, irregularly alternating; 
uterus- opening surficial, ventral, on the same level with the 
genital cloaca or very slightly behind it. Only one set of geni- 
talia in each proglottis. Testes in medu3,ll\ between the nerve 
strands. Muscular ve3icula seminalis outside of the cirrus-sac 
absent. Receptaculum 3sminis large, sharply separated from the 
apermiduct. Ovary not exactly in the median line but slightly 
approaching the margin bearing the genital cloaca, ventral, as is 
the shell-gland. Uterus in the form of a 3ac developed by the 
I enlargement inwardly of that portion of the duct passing thru the 
cortical parenchyma. Eggs without opercula. 

Type genus : Marsipometra Cooper. 

The above characterization of the. subfamily is the result 
of a comparison of M. has tat a with the existing subfamilies of the 
Diphyllobothriidae in none of which it could be located. As re- 
gards the general form of the scolex and the facts that the genital 
cloaca is marginal and that a vesicula seminalis is absent, it comes 
closest to the Triaexiophorinae; otherwise, however, it is related 
to other subfamilies. External segmentation is distinct and very 
regular, a neck being present as in Diph yllobothrium and Bothridiu m 
of the Diphyllobothriinae . The uteru3-ppening is on the same level 
with the genital cloaca, and not ahead of it as in the Tria eno- 



110 

phorinae. As in moat of the 3ubfamilies there ia only one set of 
genitalia in each proglottis. The testes are situated in the 
medulla between the nerve strands a3 in the Haplobothriinae . Un- 
like the structure in the Triaenophorinae, the receptaculum 3eminis 
is large and sharply separated from its continuation, the 3permi- 
duct, which al30 obtains for the Ligulinae, Haplobothriinae, 
Diphyllobothriinae and CyatlrCocephalus . The ovary is comparable 
to that of Tri aenop ho rus , Anchi 3tro ee;:halus and Ancn chocephalus 
(cf . Luehe, 1902a : 325) in that it is not exactly in the mediaa 
line but towards the margin bearing the genital cloaca. A3 in the 
Triaenophorinae, however, the uterus "nie die sogenannte Rosetten- 
form bildend, vor seiner Mundung mei3t etwas erweitert, ohne da33 
inde3sen diese Erweiterung verhaltnissmassig so betrachtlich ist, 
wie die sogenannte Uterushohle der meis'ten Ptychobothriiden . " This 
latter difference is further emphasized by the fact that at no 
stage in it3 development is the beginning of the uterus, which 
might be considered at first sight to be a true uterine duct, 
sharply separated from the enlarged portion as in the Ptychobothri- 
idae, which has been discussed above. The outstanding feature that 
the eggs are nonoperculate has been noted under the remarks on the 
family. 

Genus Marsioometra gen. nov . 

Generic diagnosis: Scolex unarmed, sagittate. Neck pre- 
sent, atrobila flat, ribbon- 3haped; proglottides almost rectangular, 
posterior borders only slightly projecting. Nerve strands far 
towards the margins, dorsal to the cirrus-3ac and vagina. Testes 



in two lateral fields united ahead of and behind the uteru3-3ac 
and central genital duct3. Vas deferens much coiled proximally, 
only weakly 30 close to the cirru3-3ac. Receptaculum seminis very 
long. Ovary reniform, wings tubulolobular , isthmus thick. Shell 
gland not in the middle of the genital complex but towards the 
cloaca, ahead of the ovary. Vitelline follicles numerous not in 
two lateral fields but continuous from side to 3ide in the anterior 
and posterior regions of the proglottis, situated among the body 
uiuacles. Uterus-3ac pouched, occupying the whole of the medulla 
dorsoventrally but not transversely. Uterus opening towards the 
margin bearing the genital cloaca. — ^^(.ttcov > & little 
pouch; puvjVyVs > the uterus. 

Type speoiea : Marsipom e tra hastata (Linton). 

On account of the fact that there is only one species 
known, the above generic characters have been arrived at by the 
writer much as in thejcase of Haplobothrium , with, of course, strict 
attention in a comparative way to those of the genera of the Tri- 
aenophorinae . The writer would like to call attention at this 
point to the great similarity between Marsipometra and Hapl obo- 
th rium in that each is found in an isolated genus of fishes, re- 
spectively Polyo don and Ami a which in turn are relegate.! to iso- 
lated families and orders. As suggested previously by the writer 
(1914a : 4) in dealing with Haplobothrium, the unique and general- 
ized nature of these two genera is doubtless due to the great age 
of their respective h03ts. On account of the fact that it has a 
typical bothriocephalid scolex Mar sipometra would seem to be the 



112 

younger of the two, for evidently a longer period of time mu3t have 
been required for the development of the peculiar trypanorhynchous 
scolex and method of segmentation of Haplo bothrium, if, indeed, 
both are not due to extreme degeneration, comparatively speaking. 

Mar sipometra hag tat a (Linton 
(figs. « - 50.) 
1897 Dibothrium hastatum Linton 1897 : 431-33. 

1300 Bothriotaenia hastata Ariola 1900 : 4-40. 

Specific diagnosis: With the characters of the genus. 
Medium sized cestodes up to a length of 110mm. with a maximum 
breadth of 3mm. at the middle. Scolex with terminal disc, deep 
boltxria and prominent posterior borders, 1.5-2. 8mm. in length, 
0.5-1. 8mm. in width anteriorly and 1,3-2.0 posteriorly. Subcyl- 
indrical neck, 0.8-1. 5mm. wide. First segments very short and 
wide, middle much broader than long and rectangular in outline, 
posterior ones quadrate to slightly larger than broad. Whole 
strobila much depressed. 

Cuticula 5|^in thickness, subcuticula 40-50. Calcareous 
bodies 18 x 13 pi. Longitudinal musculature weakly developed, 
that of scolex strong. Nerve strands 15 to 25 in diameter. Four 
main excretory vessels in the strobila. 

Genital cloaca 40-60 ^ in depth, at the middle of the 
margin of the proglttis; irregularly alternating; hermaphroditic 
duct present, also sphincter cloacae. Vagina opens immediately 
ahead of the cirrus. 

Testes ellipsoidal, 60-90 (jl, in diameter, 80 to 120 in 



113 



number, arranged in a 3ingle layer in the medulla and interrupted 
only centrally. Vas deferens a circular mass of coils, 0.35 to 
0.30mm. in diameter dorsal to the uterus-sac, or to one 3ide of it. 
Seminal vesicle within the cirrus-sac, 50 to 50 in diameter; 
cirrus proper slender, 0.30mm. in length, 3-15fA- in diameter. 
Cirru3-3ac elongate, flask-shaped, 0.35mm. in length, 110 ^ in 
maximum diameter. 

Vagina 15 to 30 p_ in diameter, passes to median line ven- 
trally then dorsal to the uterus. Receptaculum seminia median, 
90 in diameter. Ovary reniform tubulolobular, 0.45mm. wide and 
0.18 long; isthmus thick, ventral. Oocapt 40^ long and 18 in 
diameter. Two ventral vitelline ducts; common vitelline duct 
20 ^ in diameter. Vitelline follicles irregular in shape and 3ize, 
among the transverse and longitudinal muscles, forming a continuous 
layer around the proglottis excepting for median circular areas 
dorsally and ventrally. Shell-gland small, compact, ventral, 115 x 
55^ . Uterine duct with only a few dorsoventral coils near the 
median line. Uterus- sac circular in outline, 1.0mm. in diameter, 
divided by deep incisions into 5 to 8 pouches, filling up the whole 
medulla dorsoventrally but not laterally; openings opposite the 
genital cloaca or slightly behind its level, in young segments to- 
wards the margin bearing the genital cloaca, in graaid ones almost 
in the median line . 

Eggs, 45 x 36 |uo , non-operculate . 

Habitat: Intestine of host. 



114 



Host Locality Collector Authorltjr 

Polyodon spathula Ohio R . , Washing- E. Linton Linton 1397 : 431 



-^oiyoaon spatnuia unio k., w, 
(type host)™ ton, Pa. 



^tne pre 
paper) 



Ill.R., Beards- K.J. Van- Cooper (the pre 
town, 111. cleave sent 

L. des Allemands, H. B. Ward 
Georgia 

Miss. R. , Keokuk, * 
Iowa 

Miss. R., Fairport, ■ 
Iowa 



Type specimen : No. 4724, U.S.N.M. Coll. 

Type locality : Ohio River, Washington, Pennsylvania. 



This species was originally described by Linton "out with 
30 little attention to the internal anatomy that up to the present 
it has remained pretty much a species inquirenda et incerta 3edis, 
a3 pointed out by Luehe (18?9c : 40; 1900a : 106); altho Ariola 
(1900 : 440-441) placed it in the now obsolete genus Both riotaenia 
Railliet. 

Linton described the color of the living worms as " ... 
at first lemon-yellow; after lying inwater for a few minutes the 
bodies become colorle3s or faintly bluish translucent, while the 
heads, remained yellowish." Regarding their method of attachment 
he 3aid: "Two pits were found excavated in the mucous and sub- 
mucous layers of the pylorus near the 3pinal valve, in which the 
head3 of a number of Difcothria were inserted." The length of the 
worm was recorded by the 3ame writer a3 from 25 to 78mm. while the 
maximum breadth was 2.7mm. As shown in the table below the largest 



115 



examined "by the writer was one 110mm. in length with a maximum 
breadth of 3mm., which, however, showed the characteristic opaque 
white uterus-sac filled with eggs in only the last eight proglot- 
tides. The scolex (Figs. 23, 24, 29 and 30) and strobila are, as 
described by Linton, " ... sagittate (when at rest and contracted), 
terminated anteriorly with a button 3haped tip (the terminal disc] 
which is bluntly rounded in front and marked off from the remainder 
of the head by a slight constriction, in life angled posteriorly; 
pits (bcthria} variable in life but usually elliptical, often with 
anterior margin acuminate and sometimes with posterior margin in- 
distinct. The head is angled posteriorly both laterally Of- 
ficially] and marginally, presenting a quite characteristic appear- 
ance in the living worm. Neck subcylindrical , narrower than the 
head. The segments begin 3ome distance (5 or 3mm.) back of the 
head, as faint transverse lines. The first distinct segments are 
closely crowded much broader than long, median segments squarish 
[but 3till much broader than long"), posterior segments usually a 
little longer than broad, rectangular, apparently separating 
rather easily. ... Posterior angles of the segments slightly 
projecting. ... Outline of most of the strobilas nearly linear 
and about the same breadth as the head. All the segments were re- 
markably regular in outline, no irregularities being observed. 
While this regularity in the form of the proglottides and in their 
gradual increase in 3ize anteriorly and change of shape posteriorly 
is especially noteworthy in this species, the writer met with a few 
sasee of intercalated triangular and aberrantly subdivided segments 
in the material studied. It should be emphasized, too, that the 



116 



whole strobila including even the scolex ia conatantly much flat- 
tened dor sovent rally, which alao assists in giving the worm a 
quite diagrammatic appearance; and a3 will be aeen below thia ex- 
tend3 to the internal anatomy. Ths following table gives the 
measurements of four apecimen3, together with those by Linton for 
comparison: 





4724 


4733 


16.392 


.1 154 


16.393 


3 17.11 


Length 


60 mm 


45mm . 


76mm 


39mm . 


61mm 


110mm . 


Length of scolex 


2.75 


1.35 


1.48 


1.75 


2.34 


3.01 


Width term. disc. 


1.8 




0.60 


0.55 


0.73 


o.so 


Depth " * 






0.43 


0.31 


0.60 


0.43 


^idth at base 


2.0 


1.7 


1.35 


1.31 


1.83 


1.95 


Derth " " 





1.3 


1.16 


0.96 


0.98 


1.10 


^idth of neck 


1.1 




0.37 


0.88 


1.40 


1.52 


Depth » " 




0.4 


0.36 


0.37 


0.43 


0.68 


Length middle sega. 




0.35 


0.75 


0.46 


0.30 


0.73 


Width of same 




2.5 


2.32 


1.38 


. 3.75 


2.44 


Length post . segs . 


1.13 


0.55 


1.5 


0.85 


0.90 


1.52 


Width of same 


3.0 


3.7 


1.4 


1.10 


3.85 


2.44 


Maximum breadth 






3.38 


1.38 


3.0 


3.0 


Measured in 


water 


alcohol 


O.W. 


O.W. 


Ale . 


Formoi . 


The cuticula 


. 3 p. 


in thioJcness, 


consist a 


of two 


layers of 



equal thickness, an outer irregular and more darkly staining layer, 
which is 3loughed off in many places, and an inner, more homogeneous 
and lighter stratum between which the outermost portion of the inner 
layer shows as a dark bounding membrane. Altho only about one half 



117 

as thick on the out3ide of the scolex. and still thinner on the in- 
side of the bothria, it was net found to be modified into minute 
spines on the edges of the terminal disc nor hinder end of the 
scolex where they might be expected. Their absence also on the 
posterior borders of the proglottides (of. Haplobot hrium glo bull- 
f orme ) is not surprising since these protrude only very slightly. 
The subcuticula varies from 40 to 50 in thickness and is made up 
of narrow elongated cylindrical cells with small nuclei, the outer 
ends of which are dendritic and quite separate from each other as 
are the bodies themselves. The meshes of the parenchyma are very 
loose and open, the spaces being large and the strands of the 
cytaplasmic framework considerably narower than the small nuclei 
which are located as usual at the intersections but surrounded by 
only a limited amount of protoplasm. Linton stated that, "The 
segments contain numerous calcareous bodies, which exhibit a con- 
centric structure." In the sections made the-.vriter found them 
fairly plentiful in all part3 of the medulla and cortex and even 
among the subcuticular cell 3, elliptical or oval in outline, the 
largest having dimensions of 18 x 13 ^ . 

The musculature of thi3 species, excepting for that of the 
scolex which is well developed, is comparatively weak, no one 3erie3 
not even the longitudinal, being especially strong, altho all 
groups are prominent in that they consist of more or less isolated 
fibres quite diagrammatically arranged. Their conspicuousness is, 
in fact, amplified by the fine texture of the parenchyma. The 
frontal or transverse series do not form a compact layer closely 
applied to the inside of the longitudinal muscles but, a3 shown in 



118 

Fig. 35, a stratum of varying thickness; owing to the degree of 
separation of the fibres, especially laterally.* The myoblastic 
nuclei of many of them can be easily seen. The sagittal series are, 
however, quite prominent, equally distributed from border to border 
of the strobila and 3how their myoblastic nuclei and surrounding 
cytoplasm very clearly, reminding one of the dorsoventral muscles 
of Abothrium rugo sum (vide infra). While the fibres of both these 
series are only slightly more numerous opposite the posterior bor- 
ders of the proglottides, where they form more or less distinct 
septa, they are very well developed in the neck and anterior seg- 
ments. The longitudinal muscles form only a single layer of loosely 
arranged fascicles of irregular size in the middle and posterior 
segments, but in the neck they form a much thicker stratum, showing 
no distinct bundles and occupying the whole of the space between 
the transverse muscles and the subcuticular nuclei. Altho, as above 
mentioned, the posterior borders of the proglottides are not very 
prominent, there is a representative series of outer longitudinal 
muscles, best seen in the middle segments where they are situated 
close to the longitudinal cuticular fibres with which they are 
easily confused. Concerning the latter all that need be said is 
that they are well developed and consist of isolated fibres which 
render the two layers all the more visible. 

The musculature of the scolex is, as might be judged from 
its size and its shape, very powerful. While the longitudinal 
muscles of the neck merely enter the base of the scolex, the trans- 
verse and sagittal fibres are directly continuous with the circular 
and radial fibres, respectively, of the latter. Here, however, the 



119 

radial fibres are quite separate from the dorsoventral fibres 
with which they :are considered to be homologous, especially 
laterally where they pass from the cuticula lining the bothria to 
the sides of the scolex as in other bothriocephalids with prominent 
bothrial walls. Farther forward the sagittal muscles proper 
parsing between the bothria are 3carce, their function being taken 
over by the very numerous and closely arranged radial fibres which 
are quite as plentiful in the median line as laterally. In the 
termina disc both transverse and dorsoventral fibres are again 
prominent, while the radial ones are absent. Posteriorly the latter 
pass down along the sharp edges of the beginning of the neck. 
Frontal sections demonstrate the presence in the\edge of the terminal 
disc as well as in the posterior borders of the scolex of two 
series of longitudinal arcuate or radial fibres arranged for the 
control of these prominent ridges. These are perhaps modified 
portions cf the outer longitudinal muscles which are very numerous 
in the scolex and converge in the anterior portions of the edges 
of the walls of the bothria to become attached to the edge of the 
terminal disc at the four respective points. 

The chief nerve strands, from 15 to 25 in dorsoventral 
diameter and from 15 to 20\x, in lateral diameter, are situated far 
towards the edges of the medulla and in the median frontal plane or 
somewhat dorsally (Fig. 35). They pass dorsal to the junction be- 
tween the lateral and middle thirds of the cirrus- sac and conse- 
quently dorsal to the vagina. In the neck they are located in the 
very borders of the medullary parenchyma, but as they pass into 
the base of the scolex they approach the median line somewhat. A3 



120 

they pass on with a varying diameter towards the tip of the scolex, 
they give off a number of branches to the walls of the bothria and 
finally enlarge in the terminal disc to two ganglia, each with a 
diameter of about 50 , which give off in turn numerous large 
branches to the immediate neighborhood. Each of these ganglia is 
divided into two large trunks which, however, continue only a very 
short distance farther forward before they are joined by two com- 
missures, to their fellows of the opposite side of the scolex in such 
a way that the two branches of the ganglia on each surface of the 
sxolex are connected. In frontal sections each of these commissures 
is seen to be bowed slightly forward into the tip of the terminal 
disc and to give off further forward on each side a large branch 
which passes farther into the latter. 

The excretory system consists of a varying number of 
vessels of which four pursue a more or less constant course thruout 
the medulla of the strobila. These are found at all levels in 
transections and are separated from each other in the transverse 
direction by different distances. The outermost two, however, are 
slightly larger and have thinner walls than the innermost pair. 
They give off numerous large branches and are connected by various 
anastomoses with each other and the more peripheral vessels. In 
the neck they cannot be followed as well, while close to thejscolex 
they lose their identity and break up into a plexus of very small 
vessles which ramifies forward thruout the latter. In the posterior 
border of the scolex, however, a small branch on each side takes a 
straight course just within the nerve strand for a 3hort distance. 
Flame-cells are quite numerous and readily discernible especially 



121 



in the medullary parenchyma. In young strobilas where no segments 
have yet been lost two comparatively large excretory vessels pass 
backward to the posterior end where they unite into a small narrow 
terminal vesicle. This in a larva 12.4mm. in length was 40 y~ 
long by 10^ wide while the diameter of the excretory vessels was 
15 (a. . 

The earliest traces of the reproductive organs in the 
form of a transverse line in either half of the proglottis (the 
rudiments of the vagina, cirrus-sac and lateral portions of the 
vas deferens) appear from 4 to 10mm. from the tip of the scolex, 
while the first eggs are seen in the uterus-sacs from 25 to 35mm. 
from the same point. The development of all of the genitalia is 
very gradual and can be easily followed in good to to preparations, 
since the diagrammatic nature of the worm, above mentioned, extends 
to the reproductive system, making this species an ideal one for 
3tudy. The cirrus and vagina open into the common genital cloaca, 
which is situated in the middle of the margin of the proglottis, 
while the uterus opens on the ventral surface, not in the inadian 
line but towards the side occupied by the atrium. The cloacae 
alternate irregularly from 3ide to side, from one to ten having 
been found occupying the same margin of successive proglottides. 
The following figures represent the number of such segments before 
the genital aperture changes to the other side in specimen No. 
16.292.1 of the above table: 1, 2, 1, 8, 1, 1, 1, 2,2, 3, 3, 1, 2, 
2, 10, 1, 1, 1, 3, 3, 2, 2, 1, 4, 1, 1, 2, 2, 3, 2, 2, 3, 6, 6, 1, 
1, 1, 1, 1, 3, 2, 1, 3, 3,2 — as far forward as the rudiments 
could be conveniently traced. The genital cloaca (Figs. 27 and 28) 



133 

is elliptical in outline when viewed from the side, its longer 
diameter being directed dorsoventrally , while in transverse sec- 
tions it is squarish in outline. The dorsoventral diameter, longi- 
tudinal dimater and depth are, respectively, 70-85|^ , 40-55 |^ and 
40-60 1*- . Into the middle of the botf-O-jim of this depression opens 
the hermaphroditic duct about 60 ^ in length, into the bottom of 
which in turn opens the vagina immediately ahead of the cirrus. 
Since the cirrus proper is a long slender tube and since the exter- 
nal portion of the hermaphroditic duct is usually found quite 
tightly closed and the end of the cirrus turned around toward the 
opening of the vagina, self-impregnation would 3eem to be quite 
probable in this species. On the other hand the fact that the 
genital cloaca is so well formed and further that it is surrounded 
by a well developed sphincter and a series of muscular fibres 
radiating out into the surrounding parenchyma, as shown in Figs. 27 
and 28, argue in favor of its use in cross-fertilization. No pro- 
truded cirri was seen, however, in the material at hand. Perhaps 
both methods of fertilization occur. 

The testes are spherical to ellipsoidal in shape, their 
longest diameters being dorsoventral, while their cross-sections 
are usually circular in outline. In segments where there are as 
yet only a few eggs in the uteri their dorsoventral and transverse 
diameters are, respectively, 85-90 p_ and 60-80^. In the anterior 
and posterior ends of the proglottis, — they are not continuous 
from segment to segment but separated by the aggregations of 
sagittal and transverse muscles mentioned above as forming more or 
less complete septa, — they form a single layer situated in the 



123 

medulla in the median frontal plane, but are widely separated in 
the middle of the proglottis by the central genital organs and 
ducts, especially the uterus-3ac. Their number ranges from 80 to 
120 for each proglottis. While the wall of the testis consists of 
a very thin membrane from which nuclei protrude inwardly, the con- 
tents are such a3 to show the process of spermato-genesis with com- 
paratively great clearness. The vas deferens forms a circular 
mass of coils, 0.25 to 0.30mm, in diameter, applied like a cap to 
the dorsal side of the developing uterus-sac and thus close to the 
inner end of the cirrus-sac. When the uterus becomes gorged with 
eggs it is pushed aside somewhat but still retains similar rela- 
tions with one of the pouches of the former, located in the direc- 
tion of the genital cloaca (Fig. 26). In the mass of coils the 
duct is usually distended with spermatozoa to a diameter of 40 p. . 
It gradually narrows down to a diameter of 15 ^ before entering the 
cirrus-sac before which there is, however, no seminal vesicle. But 
within the pouch the vas deferens enlarges to form a large seminal 
vesicle, which with a diameter of from 50 to 50^, takes only a few 
coils before passing on as the cirrus proper from which it is 
3harply separated (Fig. 28). The cirrus is a slender tube from 
0.17 to 0.22mm. in length within the pouch and from 15^ in diameter 
nearest the seminal vesicle to By. at its opening. It is lined 
v/ith a thin cuticula which is circularly cleft in its proximal one- 
third but almost smooth for thejrest of its length, nowhere, however, 
showing anything in the nature of an armament. The cirrus- sac 
(Fig. 28) is an elongated flask- shaped structure with a maximum 
diameter proximally of J.10uy and distally of 40^ and a length of 



134 

0.35mm. The neck of the organ usually shows a couple of dor30- 
ventral curves, while about 20 j> of its distal end protrudes into 
the hermaphroditic duct. Its walls are comparatively thin and 
composed of an inner layer of circular muscles and an outer weaker 
and much less compact layer of longitudinal fibres. Apart from the 
seminal vesicle which occupies almost the whole of the proximal 
enlarged portion and the narrow cirrus the contents consist of only 
a limited amount of parenchymatous ti^3ue and a very few feeble 
retractor muscles. The whole structure of the cirrus-sac is in 
fact such as to suggest that the function is that of an organ for 
the expulsion of spermatozoa rather than for the emission of a 
capulatory organ; altho a few muscles passing from the body wall 
around the cloaca to the anterior part of the neck of the sac 
(Figs. 27, 28) would seem to indicate that a small portion at least 
of the cirrus is protruded, perhaps during self-fertilization. 

Altho the vagina opens into the hermaphroditic duct directly 
ahead of the cirrus, it almost immediately curves around the diatal 
portion of the cirrus-sac to the anteroventral side of the latter 
which it follows closely towards the median line. Close to the 
wall of the inner end of the cirrus-sac, however, it crosses the 
distal coils of the vas deferens towards the dorsal surface and 
skirts the uterus-sac. When it reaches the median line above the 
sack it turns sharply downward and backward. The vagina has a 
diameter of from 15 to 20 ja, opposite the middle of the cirrus-sac 
and is lined with only a comparatively thin layer of cuticula. It 
very gradually expands after crossing the inner end of the cirrus- 
sac to form a much elongated and very spacious receptaculum seminis 



135 

the diameter of which close to it3 inner end may be as much a9 90 M- • 
Ti.is is usually filled in sections with spermatozoa, a 3tream of 
which can^pften be seen passing on into the spermiduct. The 
beginning of the duct is surrounded by a poorly developed layer of 
olrcular muscles which are almost absent from the inner expanded 
portion. The receptaculum is sharply separated from the spermi- 
duct which has a diameter of only 15 to SOyu and a length of 0.12 
mm. The latter is an almost straight tube passing in the median 
line from the more dorsally situated receptaculum to it3 point of 
union with the oviduct close to the ventral wall of the medulla 
(Fig. 25). It 3hows best in transections where its walls are seen 
to be composed of an epithelium of cubical cells lying on a dis- 
tinct basement membrane, and to be surrounded with a thick layer of 
nuclei and extremely few, if any, muscle fibres. The ovary 
(Figs. 25, 26) is a somewhat kidney shaped tubulolobular organ 
situated in the posterior half of the proglottis behind the devel- 
oping uterus-3ac with it3 concavity directed forward and not ex- 
actly in the median line but slight approaching the cloaca. It 
averages 0.45mm. in width by £.18 in length. The isthmus, which is 
almost as long and about one half as thick as the wings, is located 
only slightly below the median frontal plane of the medulla. Ova 
from the same have a diameter of from 20 to 35 |x • In granid pro- 
glottides where the uteru3 is filled with eggs only a small portion 
of the degenerating ovary remains and this is accommodated between 
the two hindermost pouches of the uterus-sac. The oviduct com- 
mences in the median line anteroventral to the ovarian isthmus as 
a somewhat cylindrical oocapt 40 [a, in length by 18 in diameter and 



12S 

not sharply separated from the rest of the duct (Fig. 35). It 
passes ventrally with a diameter of 20(^ for about 50 yu before 
being joined by the spermiduct, and then for only a short distance 
farther anterolaterally along the ventral transverse musculature 
before meeting the common vitelline duct. The latter is formed by 
the union in the usual manner of two vitelline ducts coming; from 
the lateral regions of the proglottis along the ventral wall of the 
medulla. It is quite short, however, and contains in sections only 
a limited amount of yolk, its diameter being at the most only 20^ • 
The vitelline follicles (Fig. 25) are irregularly ellipsoidal in 
shape, and situated either just within the transverse muscles, be- 
tween them and the longitudinal muscles, among the latter or even 
slightly outside of the longitudinal muscles. While they vary con- 
siderably in size and, not being very numerous, are widely spaced, 
their average maximum diameter is about 50 ja, . They form a contin- 
uous band completely surrounding the medulla, excepting for irre- 
gularly circular areas above and below the central ducts and organs, 
in the median line, but are not continuous from joint to joint. 
On the whole they remind one of the vitellfcirUof A. crassum . The 
union of their . efferent ductlets can be easily traced, especially 
in frontal sections of younger proglottides, since they are com- 
paratively large and hence quite distinct. In granid proglottides 
they have, like the ovary, all but disappeared, their function 
having been almost completely performed. The shell-gland is a 
small oompaot organ, about 115 j*. in width by 55 (ul in length, sur- 
rounding the oviduct just beyond the entrance of the vitelline duct, 
or to be more exact, just beyond the first turn taken by the latter 



127 

in its return to the median line after passing laterally, as above 
stated. It is thus situated ventrally and a short distance from 
the median line. Beyond the shell-gland the oviduct continues as 
the uterine duct which makes only a few dorsoventrol coils near 
the median line "before emptying into the uterus-sac. The latter 
is formed in development by the gradual enlargement dor sally of 
that portion of the duct which traverses the cortical parenchyma 
on the ventral surface of the proglottis. Just before eggs appear 
in the sac this part of the tube can be seen in transections as a 
spindle-shaped dilatation, whose nucleated epithelial wall is sur- 
rounded by a thick layer of nuclei, the whole being, however, not 
distinctly separated from the proximal portion of the tube (the 
uterine duct of older stages) at a constriction just within the 
transverse musculature. In proglottides farther ahead this con- 
striction is outside of the transversejmuscles in the cortex, so 
that we must look upon the uterus-sac, then, as being formed by a 
gradual enlargement of the distal end of the uterus as it becomes 
filled with eggs and not as a 3ack separated in the rudiments from 
the proximal uterine tract a3 in the Ptychohothriidae . In one 
case where only 5 or 6 eggs appeared in the lumen the uteru3-sac 
had a diameter in frontal sections of SOjjl ; in the next segment 
following it was enlarged in all dir ctions, somewhat elliptical 
in outline, with a diameter of 240\l ; in the next still larger; 
and in the fourth somewhat pointed anteriorly. From then on it 
quickly enlarged until finally it is a capacious sac, as much as 
1.0mm. in diameter occupying the whole of the dorsoventral diameter 
of the medulla and almost all of the longitudinal and transverse 



128 



diameters in gravid proglottides. While as seen in transverse 
sections it is almost entire in outline, in frontal sections it is 
divided into from 5 to 8 large irregularly shaped lobes or diver- 
ticula, the hindermost two of which enclose the remainder of the 
ovary and the central connections of the reproductive ducts, as 
above mentioned. Vent rally the sack i3 funnel-shaped towards the 
small opening which only appears when the proglottis becomes quite 
gravid. Since the uterus- sacs, even the most gravid ones are not 
situated exactly in the median line cut towards the margins bearing 
the genital cloacae, the openings form " ... a zig-zag line of 
minute pores jwhichj traverses the median region of one of the 
broad faces of the strobila, each pore being near the middle of 
its segment." Linton correctly considered them to be for, the 
escape of the eggs. Anteriorly where the uteru3-sacs do not yet 
contain eggs these pores, — in reality the ventral funnel-shaped 
portions of thejsacs, — are located about 0.18mm. on each side of 
the median line, but posteriorly they are relatively much closer 
together, intact almost exactly in the median line. Furthsemore , 
they are directly opposite or slightly behind the level of the 
genital cloaca. The opening is formed only when the proglottides 
become quite gravid by the rupture of the body wall in a very 
small and limited area, but not of a preformed membrane as in the 
Ptychobothriinae . 

Concerning thejeggs Linton (p. 433) said: "The ova are 
nearly spherical, with thin shells. They are a"; out 0.04mm. in 
the greatest diameter." Those from the material preserved in 
formalin were found by the writer to be sometimes spherical in 



129 



shape but usually ovoid or ellipsoid, with maximum dimensions in 
the latter case of 45 x 36j/l . Neither in sections nor in prepara- 
tions of eggs from the uterus sacs of material in formol, alcohol 
or cleared in oil of wintergreen were opercula to be found, but at 
the one pole of the egg a small boss about 5p in diameter which 
is often enlarged to form a distinct projection or appendage. 
Alt ho development had not progressed in any of the eggs studied 
so far that the 3ix hooks of the oncosphere were visible, the 
writer is of the opinion that even in mature eggs no opercula would 
be found, 3ince its almost spherical shape and the presence of the 
button-like thickening at one pole are quite like conditions in the 
nonoperculate egg of Ab othriurn rtt°;os um, for instance, as described 
and figured by Schauinsland (1885 : 527) and further, since in 
the egg of the operculate type, as in that of D. latum or of T* 
nodulosus , described and figures by the same writer, the operculum 
is present long before the hexacanth embryo has developed. 

As regards the life-history of this species nothing is as 
yet known. It is noteworthy, however, tha+ very young larvae, such 
as shown in Fig3. 29 and30, can be easily recognized on account of 
the peculiar character of the scale n, so that it would not seem 
difficult to pick them out of the intermediate host, whatever that 
may be. All sizes from the youngest (Fig. 29) to the largest were 
present in the material studied. 

The material of this species consisted of Nos. 4734 and 
4783 U.S.N.M. Coll., Nos. 13.392, 16.431 and 17.11 C.TJ.I11 and 
No. 154 of the writer's collection, all from the intestine of 
Polyodon spat hula , the Paddle fish. 



130 



Subfamily 5, TRIAENOPHORINAE Luehe, 1839. 

Scolex armed or unarmed, always with two typical and not 
very deep bothria, ahead of which the flattened termination of the 
scolex projects more or le33 prominently in the form of a ring. 
External segmentation present or absent; in the former case an 
unjointed neck being absent. Opening of cirrus and vagina 
■arginal, irregularly alternating; uterus opening surficiaft, ven- 
tral, ahead of the marginal genital aperture. Genital apparatus 
always single in each proglottis. No muscular bulb (E3chrioht's 
body) on the inner end of ths cirrus-sac. Receptaculum seminis 
comparatively small, not always sharply separated from the narrow 
innsr end of the vagina. Uterus a much coiled canal, which while 
never forming a rosette is usually somewhat enlarged before its 
opening. 

Sexually mature in the intestines of fishes and marine 
turtles; larval conditions mostly unknown. 

Tyr e g e nu s : T r i a e nop ho r lis Eudolphi , 173 3. 



Genus 1 Triae nophorus Rudolphi, 1793. 

Ves ica ria , Cyati oe rcua et Auctorum. 
Ta¥n~i~a~ (Umn .part . ) 

Triaenophorus vel Tricus pidaria Rudolphi 1793 : 44 

Tri ouspid&ria » 1733 : 43-44 

Triaenophorua ■ " : 44 

Rhytelminthus (part.) Zeder 1800 : 317 

Rhyti s (part.) ■ 1803 : 391 



131 



Triouspidaria 


Rudolph! 


1802 : 


39-10 


n 


tt 


1809 : 


7, 35- 


illil; UO[- HO x U s 


ii 


it 




Triouspidaria 


ii 


1810 : 


32 


R 


Lamarck 


1816 : 


139 


Triaenophorus 


Rudolph! 


1819 : 


135, 








598 


N 


Creplin 


1839 : 


295 


n 


Duiardin 


1345 : 


325 


it 


Diesing 


1850 : 


304 


n 


Baird 


1853 : 


93 


tl 


Molin 


1858 : 


134 


n 


tt 


1361c: 


236 


ii 


Diesing 


1833 : 


246 


n 


Olsaon 


1867 : 


53 


it 


Loennberg 


1389 : 


41 


it 


01s3on 


1893 : 


20 


ii 


Luehe 


1899 : 


37-33 


n 


tt 


1839a: 


712 


it 


Braun 


1300 : 


1394 


it 


Luehe 


1900a: 


98 


Tri ouspidarla 


Stiles and Haasall 


1902 : 


22-34 


Triaenophorus 


Luehe 


1310 : 


23 



Generic diagnosis: Scolex armed with four three-pointed 
hooka, nevsr replaced by a p3eudoacolex . External segmentation 
completely absent. Longitudinal nervea dor3al to the cirrus-sac 
and vagina, close to the lateral borders. Testes between the 



133 

nerve strands only, filling up the whole medullary parenchyma , so 
far as this is not occupied "by other organs; a testis free middle 
field i3 quite as infrequently present aft.* pronounced dorsal layer 
of the testes. Coiling of the va3 deferens in its proximal, almost 
ttedially &it£«ated part, that portion passing distad to the cirrus- 
sac very slightly coiled. Vitelline follicles form a continuous 
mantle between the subcuticula and the longitudinal musculature, 
which i3 broken only at the places where the genital J.ucts open. 
Ovary, approaching the lateral border bearing the genital openings, 
lis s on the ventral transverse musculature, yet individual ovarian 
tubules extend partly thruout the whole medulla. Shell-gland just 
as infrequently median as the ovary, lying behind it, al30 usually 
ap/ roaching the dorsal surface somewhat. First portion of the uter- 
us only a weakly coiled canal (uterine duct) which passes thru the 
proglottis transversely and leads into a large single caity 
(uteru3-sac) which lies not exceptionally ahead of, but yet partly 
near the ovary, and usually not median but away from the margin 
bearing the genital openings. The latter al30 applies naturally 
to the uterus-opening which breaks thru later. Eggs thick- shelled, 
operculate. 

Type species: T. noduloaa (Pallas, 1781) Rud., 179 3, 

Altho, as indicated in the above synonymy and as contended 
by Stiles and Hassall (1903d : 33-24), Rudolphi should not have 
changed the name of the genus in 1819 from Triousoidaria to 
Triaenophorus again, after having U3ed it in connection with the 
specific description in 1810, the change has become 30 firmly es- 



I 



133 

tablished in the literature that it doe3 not seem perhaps justi- 
fiable to revert to the older name Tricuspida ria whioh la knov/n 
to only a comparatively small group of zoologists. 



Tria enopho rus ap . 1 a r v . 

Since all of the material at hand was larval, not even 
the earliest tracea of the reproductive rudiments showing in toto 
preparations of the largest specimens, it was, of course, impossible 
to determine the species with certainty. 

Two types of scolices were present, however, and these 
agreed with the descriptions of the organ given byjvarious authors 
for T. nodulosus (Pallas) and by 01s3on (1833 : 20-31) and Fuhrmann 
(1910 : 88-89) in particular, for T. robustus Olsson. It will be 
seen al30 in the table below that these two forms were found re- 
spectively encysted in the liver, on the visceral organs or in the 
wall of the stomach, and free in the intestine of the hosts, — or 
so firmly attached to the wall as to be deeply imbedded, the 
mucosa forming a protruding collar around the worm, — the only 
exception being those from the intestines of Esox masquinongy and 
Stizostedion vitreu m (vide infra) . Olsson pointed out that these 
pro 3peciea can be readily differentiated from each other on account 
of the situations in which they undergo their development. 'Whereas 
the larvae of T. nodulo3u s found generally within cy3ts in the 
liver of the intermediate heats, as recorded by a number of writers, 
those of T. robuatua are constantly encysted in large numbers in 
the flesh, — 01 3 son having found them in Coregonus albula and 0. 



134 

lavaretus , Luther (1909 : 53) in C. albula , and Fuhrmann in the 
"bracket" (? E30X lu ciua ) . 

The scolex of the robuatua type, shown in Figs. 31 and 32, 
is, as described by 0l3son, in the form of a truncated rectangular 
pyramid, that part immediately behind the terminal disc being con- 
siderably constricted and more nearly elliptical in transverse 
section. A3 stated by Fuhrmann, "La limite posterieure du scolex 
de T . robust us est nettement marquee et les deux bothrias, I'un 
dorsal l'autre neutral, 3on tres profonda ... 11 This delimita- 
tion of the scolex is emphasized by the fact that immediately be- 
hind the posterior border of the bothria the dorsal and ventral 
surfaces of the body of the larva are distinctly concave as are 
also the lateral 3urface3, quite diagramiiiatically, in fact, as shown 
in the figures. These concavities extend farther back for a few 
millimeters and then gradually flatten out and pass insensibly in- 
to the convexities which together form the elliptical outline of 
the cro3s-section of the middle of the larva. And the writer vvould 
like to emphasize here that thia was found to be a constant feature 
of all the material 3tudied and not simply due to any possible 
local collapsing during dehydration. Altho, as shown in the table 
the meaaurementa of the whole aoolex are much smaller than those 
given by Fuhrmann, as might be expected, it i3 chiefly the structure 
and size of the trident of hooks that led the writer to consider 
tiiia type of larva to belong to T. robu3tus. Fig. 33 of one of 
these compares very favorably with t'..ose shown in Olsson's Figs, 
31 and 32 and Fuhrmann' s Fig. 2, while the measurements (see table) 
quite agree with those given by the latter. The base of the trident 



135 

is comparatively long or deep (in the sagittal direction), hence 
the specific name according to Olsson, while not only the full 
length of the larger hooks but also a good deal of the median ones 
project thru the cuticula as the functional tips; in Fig. 33 which 
%$ from an alcoholic specimen these are seen to be darker than the 
basal piece. The following measurements are given for comparison 
7/ith Fuhrmann's of adult specimens, which are placed alongside, the 



data in parentheses be 


ing of the 


opposite trident 


on the same sur- 


face of the scolex in 


question : 












163.1 


163 . 2 


15.47 


360 . 1 


After 

Fuhrmann 


Length 


147.5mm. 


123.0mm. 


95mm . 


79mm, 


310- 370mm 


Breadth 


1.07 


1.16 


1.11 


1.01 


4-4.5 


Length of scolex 


0.93 


0.96 


1.07 


0.38 


1.14-1.3 


Breadth of term. disc. 


0.77 


0.33 


0.63 


0.74 


0.95 


Breadth scolex post- 
eriorly 


1.05 


1.07 


1.11 


1.11 


1.4-1.5 


K'idth trident 


0.31(0.30) 


0.30(0.30) 


0.26 


0.25 


0.29-0.32 


Length, mediad 


0.25 (0.24)0.23(0.33) 


0.22 


0.23 


0.34-0.28 


■ externally 


0.18(0.13) 


0.16(0.13) 


0.13 


0.13 


0.13-0.2 


Measured in 


O.W. 


O.tf . 


To to 


Alcohol 



Olsson spoke of the larva of T • robuatus being provided with a 
narrower cylindrical "cauda" as in certain Tetrarhyn cnus larvae, 
and gave the length of one as 130mm., while the anterior portion was 
approximately 60mm. long. In only a few of the larger larvae at 
hand were the remains of such a structure seen posteriorly, while 
in lot ilo.36 (vide infra) many had appendages of varying lengths 
and degrees of distinction from the fore-body, a medium large one, 



13G 

for instance, having these measurements: Length of anterior portion, 
48mm., of cauda 34, of scolex, 1.16; width of fore-body, 0.66, of 
cauda, 0.37 (3 : 1, — Olsson) . 

On the other hand, the other type of larva which was con- 
sidered oy the writer to belong to T. nodulosus , is characterized 
by a much shorter, narrower body smaller at irregular intervals, 
owing to differences of contraction, into nodules, whence the 
specific name, and by a quite different scolex provided with the 
well known form of trident (Figs. 34 to 37). While the latter and 
the scolex as seen in surficial view agree in essentials with the 
descriptions and figures given "oy various authors, e.g., Ruaolphi 
(1810 : 32-37, Tab. IX, Figs. 6-11), Wagener (1854 : 26-37, Tab. 3, 
Figs. 17-21), Olsson (1893 : 20-21, Figs. 28-32) and Fuhrmann 
(1910 : 88-89, Fig. l), it cannot be said of the material at hand 
that, a3 stated by the last writer, " ... chez T. nodulo sus on ne 
peut voir ancune limite entre le scolex et le coil du Bothrioce- 
phale ... " For inlfLateral view (Fig. 35) the "bothria are dis- 
tinctly separated from the beginning of the body, altho this is, 
as just mentioned, not nearly so apparent in 3urficial view. As 
shown in Fig. 34, the middle hook of the trident does not protrude 
thru the cuticula, since it is the root for muscular attachment, 
the upper median hook (cf . Wagener) which does protrude in the 
adult, being evidently not yet developed (Figs. 36 and 37). For a 
short distance behind the scolex the body is somewhat rectangular 
in oross-section, the sides of the rectangle being, however, slight- 
ly convex and not concave as in the robus tus type, and hence not 
30 very different from the cross-3ection of the body and farther 



137 

"back. But the material contained in lot No. 53 from the intestine 
of Egox maaqulnongy does not strictly answer this description since 
the Cody ju3t behind the scolex is slightly concave on all sides. 
Otherwise the specimens are distinctly of the no lu locus type. It 
3hould be mentioned, too, that one of this lot showed a very 3hort 
but distinct caudal piece, which with the general stout appearance 
of all of them may be accounted for by the possibility that they 
have reached the intestine of one of their final hosts, — altho 
no other specimens were taken from the M&skinonge , — and contin- 
ued their development. Likewise a few of the specimens of lots 
Nos. 10213 and N.B. 28a (vide infra) were provided with short 
caudal appendages. The smallest example of thi3 type and of all 
the material, for that matter, at hand was No. 188 of the accom- 
panying table. Altho it is only a little over two and a half 
millimeters in length, its posterior end shows that a portion, per- 
haps a caudal piece, has been torn away. The following table gives 
measurements of a number of specimens of the nodulo3Us type, similar 
to those given above for the robu stus type, with Fuhrmann's data 
for comparison: 

After 





193 


53 


151 


188 


Fuhrmann 


Length 


Piece 


Piece 


Piece 


3.68 


120-130 


Breadth at Middle 


0.61 


0.37 


0.43 


0.30 


2.5-4 


Length of scolex 


0.93 


0.63 


0.55 


0.55 


0.95 


Width term, disc. 


.43 


0.35 


0.37 


0.31 


0.37-0.47 


" scol. post'ly. 


0.64 


0.37 


0.37 


0.26 


0.57-0.6 


" trident 
■ " medially 
" " externally 
Measured in 


0.19 
0.14 
1.11 

.w. 


0.15 
0.13 
0.09 
O.W. 


0.15 
0.11 
0.08 
O.W. 


0.13 
0.14 
0.10 
O.W. 


0.135 
0.073 

0.033 



138 



It will be noticed that in spite of the fact that all of 
the measurements of the tridents are larger than those given by 
Furhmann, they are considerably smaller than those of the other 
type. 

Finally, altho no specimens of either type of larva so 
young that the hooks had not yet developed, were met with, those 
of lot Ho. 40 from the intestine of Stizofttedian vitr eum were 
provided with only very 3ma,ll tridents of the nodulosus form, the 
bases of which were not yet well developed, while the whole sco- 
lices were a sort of compromise between the two types in shape but 
of the nodulosus type as regards size, as shown in. the following 
measurements: Length, 13mm.;. width at middle, 0.34; length of 
3colex, 0.87; width of terminal disc 0.64, same of 3colex pos- 
teriorly, 0.80; width of trident, 0.14, length medially, 0.07, 
externally 0.05. Altho these ape ci email would aeem to repreaent 
an intermediate stage between the two types of scolex, so far as 
the general 3hape is concerned, the writer was inclined to consider 
them as belonging to the nodu l osus type, yet it must be said that 
smaller scolices, e.g. No. 151 of the table, have considerably 
larger hooks. 

On the whole, then, the bulk of the evidence given here 
tends to show that here in America we have probably two species, 
very closely related to if not identical with the European T. nodu- 
losus and T. robustus which have been clearly distinguished by 
Fuhrmann (1310) and also recognized by Luehe (1910 : 33). However, 
none have as yet been reported for this continent, so far aa the 
writer is aware. 



139 



The material studied is here listed as a h09t record also: 

Lot Host Loca tion Locality Collector 

Type robu s tus : 

36 Esox luoiue Intestine Flat- Rock L. Cooper 

Muakoka,Ont . 

3Sa) " 1 Go- Home R. , " 

36b) Mu3koka,0nt. 

38c " " Go -Home Bay " 

163 " ? " " ■ 

183 " " " " 

184 " ? " " " 

161 t ? " " 

163 Lota maculosa Intestine Off Giant's n 

Tomb Id. , Geor- 
gian Bay, Lake 
Huron . 

15.47 "Lake Herring" ?"" Lake Superior H.B.Ward 

Type nodu lo3us : 

71 Perca-f lavea cens In liver St. Lawrence R . , Cooper 

Iroquoi3, Ont . 

151 tEicr opterua 

dolomiea On viscera Go-Home Bay 

188 Mi c rop torus " " " 

dol omiea 

195 Catostomu3 

"* "oom;ner3onii ? Georgian Bay " 

Eh2b Notroyia deli oat U3 ? Charlevoix, Mi oh. H.E.Ward 

10213 "White Bass" Liver ? " 

U.S. 38a Stizo3tedion Stomach wall New Baltimore, ■ 
- C'-naden se Mi ch . 

40 S. vitr eum In intestine Flat-Rock L. Cooper 

53 Esox ma s qui- " " Go- Home R . " 

nongy 



140 

Family 3, PTYCHOBOTHRI IDAE Luehe, 1903, 

Scolex unarmed, with two separate and more or le33 strongly 
developed bothria or exceptionally replaced by a pseudoscolex . 
Neck absent. External segmentation never absent, but frequently 
incomplete or obliterated thru secondary foldings. Genital organs 
numerous, but only single in each proglottis. Both surfaces of 
the chain of proglottides, apart from the genital openings similar. 
Cirrus unarmed, with cleft cuticula. Opening of cirrus and vagina 
behind the uterus opening, surficial or marginal, in the first case 
on the opposite surface to the uterus-opening and almost median. 
No muscular bulb at the inner end of the cirrus- 3ac. Receptaculum 
seminis, when present, has the form of a small bind 3ack 3ituated 
at the inner end of the vagina. Ovary and sheli-glajfl median. 
Testes in two lateral fields. Uterus never taking the rosette 
shape, but usually forming a capacious undivided uterus- sac. Eggs 
thin- shelled, without oxjercula; embryonic development in the uterus 
and in consequence of exhaustic production of eggs (but dependent 
on the time of year in the case of many species) all the eggs of 
the whole tapeworto are at the same stage of development. 

Sexually mature in the intestine of fishes; larval condition 

unknown . 

In his first diagnosis of the family Luehe (lS02a : 326- 
327) emphasized the similarity of both surfaces of the strobila 
(on contradistinction to conditions in the Acanthophallidae ) , the 
unarmed cirrus with cleft cuticula, the peculiar coecal recepta- 



141 

culum seminia and the absence of opercular in the|eggs, but de- 
scribed the uterus as follows: "Uterus nie die sogenannte Po set ten- 
form annehmend, wohl aber in der Regel eine geraumige Uterushohle 
bildend, welche die ubrigen Oenitalorgane , ohne dass freilich deren 
Ruckbildung eintritt, buchstablich an die Wand drangen kann, indem 
die ganze Proglottis in reifen Proglottiden vielfach als ein ein- 
ziger sackf b'rmiger Fibehalter mit verhaltni smassig sehr dunnen 
^andungen erscheint." On account of the fact that the uterus of 
Raplobo th rium was found to answer this description in that it is 
divided into a uterus- 3ac and uterine duct, while the remaining 
reproductive organs are distinctly diphyllebothriidian in their 
nature, the writer pointed, out the difficulties in this connection 
by saying (1914a : 3) : "As a matter of fact the whole question of 
the division of the uterus into distinct regions is one concerning 
which we cannot come to any definite conclusions since, to my 
knowledge, there is no adequate description of the developmental 
relationships between the uterine tube and the uterine sac in those 
genera in which they appear." The conditions in Kaplo bothrium 
and Mars iporne tra have already been discussed above. Here will be 
given the observations on the development of the uterus to which 
reference wadjmade . 

In Bothrioc ephalus scorpi i the lumen of the uterus- sac 
appears suddenly and with a diameter of 90pu, the rudiment ahead 
showing as yet no signs of forming a cavity. This enlargement is 
situated at first, however, in "the cortical parenchyma and among the 
longitudinal muscles, only the inner tip of the structure in trans- 
sections going into the medulla. Just within the inner transverse 



142 



muscles this inner portion of the sac is joined by the uterine duct 
■ which with a diameter of 30^- elsewhere is here only 8 pu in dia- 
meter. Furthermore in the genital rudiment of the next proglottis 
ahead there is a distinct demarcation between the aggregation of 
; nuclei that will form the sac and the axial rudiment of the uterine 
duct. The 3ame separation of sac and duct with the narrowing of 
the latter just before entering the former is present in the follow- 
ing segments even where the first eggs are to be seen in the lumen. 
Thus the eggs must have passed this narrowed region which is a 
great deal smaller than their diameters. Still farhter back where 
the lumen is about 165 ^ in diameter there can be seen not only the 
situation of the sac in the cortex and among the longitudinal 
muscles, projecting as yet only a short distance into the medulla, 
— altho here the bundles of muscles are deflected peripherally, — 
but also the separation of the two parts by a narrow neck only 10 ^. 
in diameter. B. cuspida tus 3hows the same distinct separation of 
the uterine duct and uterus-sac in the proglottides where there 
i are already a few eggs in the latter. In Clestobothriu m crassiceps 
conditions were found to be quite the same. When the lumen of the 
sac attains dimensions of about 60 x 35 ju, and is lined with an 
epithelium which takes the conter stain more like a cuticula but 
shows distinct nuclei on its surface towards the lumen, the uterine 
duct opens into it with a distinct reduction in diameter. The 
epithelia of the two are, however, quite similar and continuous, 
the nuclei being located in a similar manner in both. Proglottides 
ahead show that the sac is formed by an enlargement of the end of 
the duct, which takes place first in that region passing thru the 



143 

cortex quite as in Bothr ioceph alus . 

Thus we see that the uterus sac of this family is quite 
different from the functional enlargement of the uterus of the 
Diphyi'iobothriiciae , with the exception of that of Kapl obo thr i um , 
3ince at all stages in its development it is sharply separated 
from the uterine duct. But a3 it was not so much this exact separ- 
ation of the two portions 5.3 the constant presence of a "Uterus- 
hohle" in this family and its absance in the other, where the 
"Rosettenform" is more common, that was emphasized by Luehe, and 

since the structure in Haplobothr lum is distinctly ptychobothrii- 
dian in character, the functional enlargement of the uterus can not 
now be considered to be of such systematic importance as was 
formerly believed to be the ea$e . 

Subfamily X. PTYCH030 THR 1 1 NAE Luehe, 1899, 

Scolex with two surficial sucking grooves, which may be 
modified by considerable growth together of their free edges. 
Genital openings surficial, those of the cirrus and vagina dorsal, 
that of the uterus ventral and ahead of the other two. Va3 deferens 
strongly coiled, dorsal. Ovary ventral; shell gland dorsal. 
Vitelline follicles usually in two lateral fields in the cortical 
or medullary parenchyma. Testes completely filling the medulla, 
mostly marginal to the longitudinal nerves which are well towards 
the median line . 

Occurrence : Exclusively in fishes. 

Type genus : Bothriocephalua (Rud.) Luehe. 



144 

The aboi-e diagnosis lack3 the wor ,ds "seldom armed" 
after n 3colex" which appear in Luehe • s latest (1910 : 24) charac- 
terization not only of this subfamily but of the family, because 
they do not appear in his earlier (18S3 : 41 and 1902a : 336, 
respectively) papers, nor doe3 there seem to the writer to be any 
occasion for their use. 





145 


Genus 1. Bothriocephalic Rud . 1808 . e . p. Luehe 1899, e. p. 


Taenia (part) 


Auctorum. 


Rhvteiminthes (part.) 


Zeder, 1800. 


Alvselminthus (part.) 


Zeder, 1800. 


Rhvtis (part.) 


Zeder, 1803. 


BothrioceDhalus (part. 


)Rudolphi, 1809. 


Bothriocephalus (part. ) 


Rudolphi, 1819. 


Pibothrius (part.) 


Rudolphi, 1319. 


Bothriocephalus (part .$> 


Leuckart, 1819. 


Bothriocephalus ( part . ) 


Dujardin, 1845. 


Bftbothrium ( part . ) 


Diesing, 1850. 


Bothriocephalus (part. ) 


Baird, 1853. 


Dibothriura (part.) 


MClin, 1861. 


Dibothrium (part.) 


Diesing, 1865. 


Bothriocephalus ( part . ) 


Carus, 1885. 


Bothriocephalus ( part . ) 


xviatz, 1891. 


Bothriocephalus (part. ) 


Ariola, 1896. 


Bothriocephalus s.str. 


Luehe, 18 99. 


Bothriocephalus (part . I 


Ariola, i«*OQ 


Bothriocephalus s.str. 


Braun, 1900. 


Bothriocephalus s.str. 


Luehe, 1910. 


Generic diagnosis; Scolex elongated, with tv;o only weakly 


developed sucking grooves. Neck absent. External segmentation 


well developed; between two consecutive genital segments there is 


always present a sav.-tboth notching of the lateral border, yet a 


corresponding transverse furrow 


on both surfaces is sometimes 


lacking. Vitelline follicles in the cortical parenchyma, contin- 


uous from proglottis to proglottis, as are the testes. Recepta- 



J 













culuro 


serainis absent. Beginning of the uteru 


s a v. 


inding canal 


(Uterine 


duct) which opens into 


a large nearly 


spherical cavity 


(uterus 


-sac or uterus s.str.). 


Uterus-opening 


approximately me- 


dian, a 


s is the dorsal genital 


opening. 






Type species: Bothriocephalus scorpii (Mueller). 


Species 1. Bothriocephalu 


s scorpii (Mueller r 1776 ) 




(Figs. 38 


- 49.) 






1722 


Vermis rauxt imembris 
rhomb i 


Leeuwenhoek 


1722 


; 402. 


1776 


Taenia scorpii 


Mue Her 


l <~i n u 


Ota 


1780 


Taenia scorpii 


Mueller 


1780 


: 17 9. 


1780 


Taenia scoroii (oart.) 


Fabricius 


1780 


: 319. 


1786 


Taenia scorpii 


Batsch 


1786 


: 235. 


1788 


Taenia scorpii 


Mueller 


1788 


: 5-6. 


1788 


Taenia scorpii 


Schrank 


1788 


: 48. 


17 90 


Taenia scorpii 


ttmelin 


1790 


; 3078. 


1799 


Taenia SfiOTTiii 


Rat ke 


17 99 


: 68. 


1800 


Alyselminthus bipuncta 
tus 


Zeder 


1800 




1802 


Taenia nunctata 


Rudolphi 


1802 


: 109-110. 


1802 


Taenia scorpii 


Bosc 


1802 


: 307. 


1803 


Rhvtis bipunctata 


Zeder 


1803 


: 296. 


1810 


Bothriocephalus puncta 
tus 


- Rudolphi 


1810 


; 50. 


18*9 


Bothrioc. punctatus 


Rudolphi 


1819 


: 138. 


1819 


Bothrioc. punctatus 


Leuckart 


1819 


: 40* 


1844 


Bothrioc. minctatus 


Bellingham 


1844 


: 254. 


1845 


Bothrioc. punctatus 


Dujardin 


1845 


: 617. 


1850 


Bothrioc. punctatus 


Beneden 


1850 


: 160. 











147 


18 50 


jPlbothriurn punc t a turn 


Diesing 


1850 : 


593. 


1853 


Bothrioc. punctatus 


Baird 


1853 ; 


89. 


1855 


Dibothrium punctatura 


Leidy 


1855 ; 


444. 


1856 


Dibothrium Dunctatum 


Leidy 


1856 : 


46. 


1858 


Bothrioc. punctatus 


Cobbold 


1858 ; 


157. 


1858 


Dibothrium punctatura 


Molim 


1858 : 


134. 


1861 


Dibothrium punctatum 


Molin 


1861 : 


235. 


1863 


Dibothrium ounctatum 


Diesing 


1863 : 


240. 


1867 


Bothrioc. punctatus 


Olsson 


1867 : 


14,55. 


1878 


Bothrioc. punctatus 


Linstow 


1878 : 


237. 


1885 


Bothrioc. punctatus 


Carus 


1885 ; 


120. 


1889 


Bothrioc. punctatus 
forma bubal id is 


Loennberg 


1889 : 


32. 


1890 


Dibothrium punctatura 


Linton 


1890 : 


731. 


1891 


Bothrioc. punctatus 


Loennberg 


1891 : 


51. 


1892 


Bothrioc. punctatus 


Matz 


1892 : 


105. 


1893 


Bothrioc. ounctatus 
forma motellae 


Loennberg 


1893 ; 


13. 


1893 


Bothrioc. punctatus 


Olsson 


1893 : 


16. 


1897 


Dibothrium punctatura 


Linton 


1897 : 


430. 


1899 


Bothrioc. biounctatus 


Luehe 


1899 : 


43. 


1900 


B.othri.oc. punctatus 


Ariola 


1900 : 


394. 


1900 


Bothrioc. bipunctatus 


Braun 


1900 : 


1691. 


1902 


Bothriop. bipunctatus 


Fuhrma nn 


3.902 ; 


446. 


1902 


Bothrioc- -ounctatus 
forma punctatus v$l 
_typ.ica 


Schneider 


1902a: 


14. 


1902 


Bothrioc nun ct.a 1 1 m 

forma rhombi 


ui^iUlC iUcI 


i q n 9 o • 


■LO . 


1902 


Bothrioc. punctatus 
forma cott i-quadri- 
cornis mihi 


Schneider 


1903 : 


75-76. 



148 



1910 Bothrioc . bipu nctatus Luehe 1910 : 25. 

Specific diagnosis: With the characters of the genus. Large 
cestodes, up tp 950mm. long by 6ram. wide. Scolex, large, elongate, 
with prominent terminal disc, widest anteriorly; length 1.0-3. 5mm. 
breadth 0.3-0.5. Bothria long and narrow, shallow posterior^. 
First segments subcuneate with weakly prominent posterior borders, 
longer than broad, middle and posterior segments much depressed, 
former very short and broad, latter relatively less so and grouped 
in twos or threes; lateral borders crenulate. Ripe proglottides 
2-4mm. wide by 0.2-0.8 long. Strobila usually incomplete poster- 
iorly. 

Cuticula 5(^ in thickness. Calcareous bodies 13pL in diameter. 
Inner longitudinal muscles in fascicles. Six chief longitudinal 
excretory vessels. 

Opening of the genital cloaca at the bottom of a dorsal medi- 
an longitudinal depression ruuning thmiout mature segments, on a 
low papilla in each proglottis and halfl way betweefa the anterior 
and posterior borders. Vaginal opening immediately behind that of 
cirrus. Distinct ductus hermaphrodit icus present. 

Testes subspherical, 35-70f>- in diameter and 30 to 60 in num- 
ber for each segment. Vas deferns a compact mass lateral to cir- 
rus-sac and opposite the uterine tube, 0.18 x O.lOmra. Cirrus-sac 
at right angles to dorsal surface, 115 x 120 x 75-80 , extending 
only a short distance into the medulla; thick layer of nuclei 

v ithin its wall. Cirrus proper, notprotruded, 65 x 15\x . 

Ovary compact, tubulolobular , 0.33mm. wide by 0.15 long 
where uterus-sac is not greatly distended. Isthmus onlyventral. 
Odcapt 35^ in diameter. Vitelline follicles in two lateral weak- 



149 

ly united fields on each surface, 350 to 540 in number, 35-55^ in 
diameter; vitelline reservoir small. Shell-gland large, 115^ wide 
by 0.85 deep, median, close behind cirrus-sac. Uterine duct vfcdju- 
minous on both sides of the median line, closely applied to ovary 
behind. Uterus-sac spherical to flattened anteroposterior^, Oc- 
cupies one-sixth of transverse diarater of proglottis, alternating 
irregularly from side to side, or often quite median. Opening in 
middle of sac, ventral and well forward, formed by the rupture 
of a distinct membrane. 

E £g s > 66-80JA- in length by 43-45 in diameter, without opercu- 
la, forming dark brown maculations in ripe proglottides as they 
show thru the walls of the uterus -sacs. 

Habitat: In the intestine of the host. 



Host 
Cottus scor pius 



Localit y 
Denmark 
Gryphswald 



Ireland 

"Oresund e 
Berg, Sweden 

Norway 

Sweden 



Arctis Ocean 

Gulf of Fin- 
land 



Collector 

Mueller 

Rudolphi 

Kais. -konig, 
nat» lkab. 

Bellingham 

Olsson 

Loennberg 
ti 



Grafverna and Olsson 
Saset,Sea of 
Bahusia 



Schneider 



Authorit y 
Mueller 1784:6 
Rudolphi 1819:139 
Leuckart 1819:40 



Bellingham 

1844:2 54 

Olsson 1867:55 



Loennbergl8 90 : 22 
" 1891:51 
Olsson 1893:16 

Linstow 1901:281 
Schneider 1902:15 



Cot tus scorpius 



bubalis 



wiurraan-Kuste 

k 

W^ite Sea 

North Sea 

Firth of Clyde, 
Millport , Scot- 
land. 

England 

Norway 

Sweden 



Coll.2iOol.Mus 
K.Akad.Wiss. 
Petrograd . 

Danilevskij 

Nicoll 



Cobbold 
Loennberg 



" Grafverna and 01s son 

Naset 

" " "Oresund e " 

Berg" 

■ " North Sea Mcoil 

" auadricornis Gulf of Finland Schneider 

Naples Rudolphi 



Pleuronectes bos 
cius 

Pleuron . f lesus 

" Jliajujaus 



"Oresund e Berg" Olsson 
Denmark Mueller 



"Turbot" 
Pleuron . rhombus 



Gryphsv aid 
Ariminus 



Ireland 

Langrunne, 
Rennes 

Belgium 

Naples 

Ireland 



Rudolphi 



Kais.-konig 
nat * lkab. 

Belxingham 



Dujardin 

Beneden 

Rudolphi 

Druramnnd 



150 

Linstow 1903:19 



Nicoll 1907:70 
" 1910:355 

Cobbold 1858:157 
LoennberglS 90:22 
" 1891:51 
Olsson 1893:16 

Ariola 1900:396 

Nicoll 1907:71 
Schneiderl903: 75 
Rudolphi 1819:139 

Ariola 1900:396 
Mueller 1784: 6 
Rudolphi 1819:139 

M MM 

.ueuckart 1819:40 



Beliing- 
ham 1844:254 

Dujardin 1845:618 



Beneden 1850:161 
Rudolphi 1819:13 9 
Belling 



ham 



JL8«4:254 











151 


Pleuron. solea 
Jo^pedo narce 


Naples 


Kais.-koni^.- 

Hal IKuD 

Rudoiphi 


Leuckart 
Ariala 


1819:40 
1900:396 




ti 


M 


Rudoiphi 


1819:139 


" oculata 




Volz 


Volz 


1900:55 


Gadus aeRllfinus 


Arctic Ocean 




Linstov/ 


1901:281 


■ minutus 


Naples 


Rudoiphi 


Rudoiphi 


1819:139 


ArnogTBSSus boscii 


ii 


ii 


Ariola 


1900:396 


" pe^osa 


Ariminus 


ii 


Rudoiphi 


1819:139 


M soiea 




ivius.Vienn. 


ii 


ii •• 


Trivia adriatica 




C.E.V. 


ii 


ii it 


U If 


Hafnia 


Eschricht 


Diesing 


1850:594 


" lineata 


Ireland 


Drummond 


Diesing 


1850:594 


roc lid ujci A-L met 


5-iHgxd. no. 


Brit .Mus # 




lo do : o y 


Platessa plana 


Pennsylvania 


Leidy 


Leidy 


1855:444 


» flesus 


Germany 




Luehe 


1910:25 


" passer 


Trieste 




Stossich 


1898:116 


Rhombus maximus 


Italy 


Molin 


Molin 


1858:134 


ti h 


Patavia 


it 


ii 


1861 :235 


it it 


"Oresund e Berg" 01s son 


Olsson 


1867 : 55 


it ti 


Trieste 


Stossich 


Carus 


1885 :120 


it it 


Venice 


Ninni 


Stossich 


1890:7 


11 N 


\warnemunde 




Matz 


1892:105 


li ii 


Rossitten, 
C r an z, kernel 






lo yo * oo 


it ii 


Trieste 




Stossich 


3.898:116 


it ii 


G8n0va 


Parona and 
Ariola 


Ariola 


1900:395 


ii ii 


Trieste 


Stossich 


Stossich 


1901:97 


ii it 


Gulf of Fin- 
land 


Schneider 


Schneider 


1902;15 











152 


Rhombus maxiraus 


North Sea 


Nicoll 


Nicoll 


1907:72 


M barbue 




Volz 


Volz 


1900:55 


II ] q "i O 


UI LcilU. w 

- Berg" 


D 1 c c r^r** 
ID X O o \JL\ 


Is s on 


loo / : oo 


ii ii 


\J 1 cx i v ci lid cliiu. 

Naset 


It 


it 


XO J7«J a ID 


" maeoticus 


Odessa 


Nordmann 


Lins tow 


1901:281 


J_l u UI 1 U p o C U L O IUCi Li. 

JLaia 


MaytVia's Vinp- 

jvicij. ilia v x 1 1 v> 

yard, Mass. 


T»1 nt rm 


Xj XI l u» Ui 1 


xc yu • / oc 


Bothus aaculatus 


Woods Hole 


Linton 


Ariola 


1900:396 


It II 


it it 


it 


Linton 


1897:430 


He mi t r i p t © r,us 




Nat .Mus. 


it 


ii ii 


ii 


Casco Bay, Me. 


ii 


ii 


it it 




Woods Hole 


ii 


if 


it ii 


it 




UOll.Wal .MUS . ) 

Wash. 


Ar ioxa 




Limanda ferrusinea Block Id. 


U.S. Fish Com. 


Linton 


1897:430 


ii it 


WoodsHole 


Williams 


it 


1901:284 


it n 


ii ii 


Linton 


Linton 


1901:485 


Labrus raaculatus 


"Qresund e 
Berg" 


Olsson 


Ar i ola 


1900 -396 


Motella raustela 




Coll. Mus. Vien. 


it 


It It 


Mullus barbatus 


Genova 


Parona 


it 


It H 


Solea monochii 




Coll. Mus. Vienn. " 


II It 


Acipenser ruthen- 

m 




Volz 


Volz 




Scorpaena porcus, 




it 


ii 


II II 


Paralichthvs ob- 


tooods Hole 


Linton 


Linton 


1901 '484 


Lota vulgaris 


Dvina-Fluss 


Danilevskij 


Linst ow 


1903:19 


Raja clavata 


Black Sea 


Pilat 


Pilat 


1906:191 


Ansuilla vulgaris River Dee. Scot 
land 


- Scott 


Scott 


1909:79 



153 

Decapterus puncta - Woods Hole Reg- - Sumner, Osburn and 

IBS ion Cole 1913:586 

Hippoalossus hippo - " •« 

^lossus 

;.i voxocepha lus aeneus " " '» 

dec iftspinosus octo - " " 

decimspi,nosus 

Palinurichth ys perci,- " " " 

f ormis 

Paralichth ys dentatus " " " " 

P^^iOsromsJ^s " 

americanus 

Scomber scombrus " " " " 

Trachuro os crumeno - " 

phthalmus 

Uroohvcis chus s " - " " " 

Hemitripterus america - Passaraa- Cooper Cooper 

nus quoddy Bay, (the present paper [ 

New Brunswick 

" " Brandy Cove, " * 

St. Croix R. ,N.B. 

" " Ifcoods Hole V.iUEdv.ards " 

? jid yoxacephalus aeneus " " " " 

? ■ groelandicus " 5 " " 



154 

As shown in the table of measurements given below most of 
the specimens of this species ranged in length from 50 to 340mm., 
while none were considered to be complete posteriorly. The smalles 
measured 28mm. in length and the largest 677mm. The 3colex 
assumes a variety of forms in preserved material, but agrees in 
general with the descriptions of the organ for B. scorpii by all 
the authors from the time of Rudolph! (1810 : 51). Its commonest 
shape is shown in Figs. 38 and 39 where it ia seen to ne quite 
elongated, somewhat broader and truncated anteriorly and narrow 
posteriorly. The anterior portion is in reality in the 'form of a 
low pyramid, compaiable internally (vide infra) as well as extern- 
ally with the terminal di3c of the Triaenophorinae . Its base is 
deeply indented dorsoventrally , that is opposite to botKrria, but 
rounded laterally. The whole scolex is broadest at about its 
middle and narrowest at its posterior end, a portion of which i3 
here considered to be ft beginnings of the lirst segment on account 
of its being in all the material at hand set off from the re3t of 
the 3Colex ahead by a more or less definite groove. The bothrium 
is in the form of an elongated V, being ordinarily widest and 
deepest just behind the terminal disc and much narower and shallow- 
er posteriorly where it is not bounded by a definite wall but 
spreads out on the base of the scolex. In many specimens, however 
the soolex is 30 contracted and the wall 3 of the bothria 30 pro- 
truded that the latter shows its greatest depth at the middle of 
the scolex. In lateral view (Fig. 39) the 3colex is more oval 
in outline and a little wider towards the base. From this fact it 
is conceivable that Mueller's (1784a) Fig. 7 showing a more 



155 



"orbicular" scolex in B. aoorpli in lateral view may be explained 
by supposing that he was dealing with a much contracted specimen, 
altho in justice to the other side of the question, it must be 
said that the first segments in his figure are by no means con- 
tracted. It will be noticed that the figures of the 3colex given 
here agree very closely with that of Scott (1902 : Fig. 3, PI. V) , 
altho from the fact that he records B. scorp ii a3 having been 
found in Angui lla vulgaris > it is quite possible that he had in 
reality B. claviceps (G-oeze) which has been found only in eels up 
to the present, so far as the writer is aware. 

Segmentation begins immediately behind the scolex, 30 that 
there is nc true neck, altho the base of the scolex has the 
appearance of a very short neck region from which the foremost 
segments are cut off as soon as they form such in fact being con- 
sidered to be the case. The anterior part of the strobila, however, 
serves the purpose of a neck in that it shows a division into sub- 
segments in a manner to be presently described. As regards the 
habit of the whole strobila and the general shape of the segments, 
Rudolphi's (1810 : 51-52) description of the species is so appli- 
cable to this form that it is given verbatim: 

" Co H um nullum. Corpus planum, margine 
crenato . Art i culi capiti proximi plerumque long- 
issimi, augU3tissimi , subcaueati , margine postico 
untrinque parum exstante, saepe tamen, praesertim 
post mortem, contract! , ut reliquis vix longiores 
apparent. Articuli insequentes anticis breviores 
et sensim latiores; postici subsequales, fere 



156 

quadrat i , utplurimum latiores quam longi, internum 
quasi ex duobus tribresve confusis oompo&i ti , satis 
magni , margine obtusiusculo hinc inde inciso. Ar- 
ticulu3 ultimus obtusus. 

Line a utrinque longi tudinalis articulos majores 
percurrit. Inter utramque faeturae apparatus." 

Leidy (1855 : 444) described the strobila of the B. scorpii 
which he found in Platesa plana as follows : "Neck none. Anterior 
segments cameate or triangular; posterior one3 quadrate; each 
with an appearance of three subdivisions, with the subsegments 
having a pair of generative apertures, in the course of a longi- 
tudinally depressed dark colored line, passing the length of the 
body,"; which Cabbold (1858 : 157) said of individuals from 
Cotfr u3 , buba l is : "Toward the lower part of the so-called neck, the 
joints exhibited at the lateral margins indications of division, 
which became gradually more defined towards the tail," In the 
same connection Krabbe (1865 : 37) said that, according to Esch- 
richt, "Pendant leur developpement ulte'rieur, 1 'augmentation du 
nombre des articles n'est pas tonjours exclusivement due, comme 
chez les Taenias, \ la formation de nouveaux articles engendres 
par la tete, mai3 chez quelques especes, telle3 que les B. dubiu s , 
variabilis et fasci atus (here B. punctatus also) elle est encore 
produite par la division transversale qui s'opere dans les articles 
deja forme's." Olsson (1837 : 55) also referred to multiplication 
of segments by transverse division of older one3. Loennberg 
(1891 : 53) denied this statement of Olsson' s, but as pointed out 
below the negation is applicable to the posterior mature segments 



157 



of this form at least, not to the middle segments referred to by 
the latter. Linton (1890 :773) said: "Secondary segments appear 
at about the twelfth segment from the head. These are formedfby 
a division of each segment into two by means of a median .transverse 
line. This is repeated farther back in much the same manner as 
described under D, miorooephalum . " In this form such subdivision 
of segments to form daughter segments was found to occur all along 
the strobila from close behind the 3colex to well into the region 
showing the median row of reproductive rudiments, and in such a 
manner that, in the anterior part of the strobila at least, what 
was considered to be a primary segment, situated between the most 
prominent transverse furrows visible, became subdivided into 2, 4, 
8, 16 and finally 33 divisions, ea,ch of the latter accommodating 
two reproductive rudiments. But it must be emphasized that this 
method of formation of new segments is only approximately followed 
out since, as it passes backward in development, the primary seg- 
ment does not always contain 64 genital rudiments, because in the 
first place some secondary or even tertiary transverse furrows be- 
come almost a3 prominent as the primary ones, and secondly, because 
at the 3ame time there is considerable further subdivision not only 
of the- peripheral tissues but especially of the rudiments them- 
selves. Close behind the acolex the primary segments are very short 
(Fig. 38), the first six to ten being divided only into two sub- 
divisions in strobilae of moderate size, but into three or four 
subdivisions in the largest chains. Farther badk this more or less 
regular manner of segmentation takes place gradually, but division 
is usually seen to occur more readily and quickly in the anterior 



158 

part of the primary|segment or of its major subdivisions, i.e., 
secondary or tertiary, than in its posterior part, — sometimes, 
however, the reverse being the case. Thus in general there is a 
sort of dominance of the anterior end of the segment, which one 
might call a zooid in the sense in which Child uses the word, over 
ite posterior end as regards metamerism. While this method of 
formation of segments i3 further obscured by the fact that often 
one seems intercalated among primary segments, showing these 
features well, others which seem to lag behind in division and 
are hence younger, and that in much elongated strobilas it is 
still more difficult to dintinguish between primary and secondary 
transverse furrows, owing to their being quite smoothed out es- 
pecially medially, the whole plan is sufficiently clear to warrant 
its being described with the definiteness here given. Fig3. 40, 
41 and 43 will give a bettei idea perhaps of the whole method of 
segmentation than this description. While in Fig. 40 the primary 
segments are indicated by asterisks, in Figs. 41 and 43 the whole 
drawing is in each case that of a primary segment. Under the 
heading of the reproductive system below it will be seen that in 
the mature portion of the strobila the most prominent transverse 
furrows are described as coming approximately every eighth or six- 
teenth genital segment. This is due to the fact that the secondary 
and tertiary furrows, respectively those dividing the primary seg- 
ments into two then four parts, become quite as pronounced as the 
primary ones, thus making it impossible to follow this system of 
segmentation beyond the region of differentiation of the genital 
rudiments . 



159 

At least three prominent longitudinal grooves run thruout 
the median and posterior portions of the 3trobila on each surface, 
even cutting thru the posterior borders in many places, but their 
course is not regular; they are accompanied by numerous other 
shorter and more irregular grooves; all of which, however, do not 
seem to be due simply to lateral contraction of the segments. The 
following table gives a list of measurements of representative 



specimens in alcohol 


from the 


materi 


al at 


hand. 








Numb pt of sr; p r i wpt\ 


196 .1 


196 .2 


196 . 3 


196 .4 


197 


198 ,1* 


198 .2 


Total length 


234 


173 


103 


51 


677 


233 


180 


Length of 3colex 


1.20 


1.07 


0.81 


1.14 


1.11 


5.00 


0.85 


Breadth of terminal 


disc 0.30 


0.33 


0.26 


0.33 


0.28 


0.29 


0.27 


Breadth at middle 


0.35 


0.32 


0.30 


0.37 


0.31 


0.11 


0.23 


Breadth at base 


0.38 


0.27 


0.24 


0.35 


0.26 


0.11 


0.16 


Length of Seg. I 


0.14 


0.16 


0.09 


0.18 


0.33 


1.83 


0.44 


Length of Post. Seg. 


0.85 


0.45 


0.35? 


0.57 


? 


0.54? 


0.81 


Breadth of same 


1.83 


1.12 


0.53? 


0.32 


? 


1.70 


0.99 


Maximum breadth of 
strobila 


2.50 


1.60 


1.05 


1.57 


3.35 


2.80 


1.51 



♦Stretched during- fixation. 



Since Loennberg (1891 : 52) described the cuticula of the 
species there has been no mention of this tissue in the literature 
so far as the writer is aware. In sections it was found to be 5a 
in thickness and composed of two lay:rs, the outer of which is 
about two-fifths of the whole thickness and is made up of rather 
3tout, closely set "cirri" which 3tain njuch more readily than does 



160 

the inner more homogeneous and lighter layer. These cirri seem 
to lie on a distinct membrane since their proximal (central) ends 
are all even and distinguishable in 3ome places as dark granules. 
In sections stained more deeply than those which show the inner 
layer as a 3ingle homogeneous stratum, the latter is divided into 
two layers, the outer of which is less deeply stained than the 
inner and about one-half as thick or one-fifth of the thickne33 of 
the whole cuticula. The wavy nature of the cuticula and the base- 
ment membrane were, found to be as described by Loennberg, but in 
many places the membrane is- separated from the cuticular muscula- 
ture by a very thin clear 3pace barely distinguisheable with high 
powers. The cuticula covering the 3colex was found to be about 
4f*- thick, the difference between it and that over the proglottides 
being due to a thinner homogeneous stratum. The outer layer of 
the cuticula is not modified to form spinelets on the posterior 
borders of the proglottides, as in C. orassioeps (vide infra), nor 
on the edges of the terminal disc, but the pseudocilia are somewhat 
longer and relatively 3touter on the scolex and anterior segments 
than elsewhere. 

The subcuticula, from 25 to 30^ in thicknes3, has the 
nuclei of its spindle-shaped cells arranged at various levels so 
that the space between the cuticula and the vitelline follicles is, 
excepting for its outer one-third, well filled with them (Fig. 43). 

The chalk-bodies, described by Loennberg were not studied 
in living material, but spherical spaces which were occupied by 
them bsfore they were dissolved out by the acetic acid of the fixing 
agent, were found to be more numerous, as Loennberg stated, in the 



161 

cortical than in the medullary parenchyma. In the scolex they 
are somewhat more numerous than in the 3trobila, in "both of which 
locations they attain a diameter of 13^. 

Loennberg (1331 : 53) gave a good description of the mus- 
culature of the spcies, while Luehe (1897a : 747) referred to that 
of the posterior border of the proglottis in the discussion of the 
arrangement of the muscles of the whole order. In addition to 
corroborating the findings of these authors it was noticed that 
the fibres of the transverse series are mostly confined to the very 
ghort regions between the sets of reproductive organs and are most 
numerous just ahead of the prominent segmental furrows mentioned 
above, this applying to both the inner and outer lots. Towards 
the median line ach layer of longitudinal muscles is about 35 |a. 
in thickness and comrosed of bundles of various sizes in which the 
fibres are very closely arranged. The outer longitudinal muscles, 
the extension of which into the posterior borders of the segments 
immediately behind the scolex are only weakly developed, are con- 
fined in the scolex almost completely to very thin bands situated 
close to the cuticular musculature in the edges of the bothria, as 
described elsewhere by the writer (1914b : 92) for H. globuli forme . 
There is also a weak series of longitudinally arcuate fibres ar- 
ranged around the edges of the terminal disc as in the latter. 

Loennberg (1391 : 54-53) described the nervous system so 
well that little needs to be added. It was noticed that the fore- 
most four large branches from the brain complex were not relatively 
as large as those shown in Loennberg' s Fig. la, and that the com- 
missure appeared to be divided into two not distinctly separated 



163 

frontal strands, the whole depth of which, including the spaoe 
between them, was not as much a3 that shown in his Fig. lc. In 
the strobila the chief nerve strands, each from 15 to 35^ in 
diameter, were found to be 3ituated towards the ventral 3ide of the 
medulla and at the junctions of the lateral and median quarters of 
the latter, as shown in Fig. 43. 

The excretory system of B. scor pii was described in detail 
by Fraipont (1831 : 3-13), while Loennberg (1891 :53-54) added some 
further notes on its structure, the former, however, working on 
living material in which the canals are much more readily seen. 
In good toto preparations the "canaux descendants" may be easily 
seen in segments showing the reproductive rudiments as well as 
farther forward. Owing to a mere accident, temporary preparations 
were made with more or less constant success, showing the details 
of the reticulum of descending canals in great detail. When some 
pieces of a strobila were being transferred from synthetic oil of 
wintergreen to a slide for the preparation of toto mounts by the 
further addition of xyiol-dam^r , they suddenly became opaque white 
and remained so for some time after the damar and cover-glass had 
been added. This opacity was found to be due to air having been 
drawn into the excretory canal3 not only thru their cut ends but 
thru the foramina secundaria. But 3ince the superficial reticulum 
and all the finer canals were filled with air, nothing of the ar- 
rangement of the larger canals could be made out until a short time 
had elapsed or until the preparation had been heated slightly. 
Then the air in the smaller canals became replaced by the xylol- 
damar or become dissolved in the latter, and the larger-canals 



163 

stood out as very distinct silver threads. This sort of pre- 
paration is unfortunately not permanent, 3ince after a few minutes 
all of the canals iisappear, excepting the largest which can still 
be followed as in ordinary toto mounts. The results of this some- 
vhat uncertain method of demonstrating theexcretory canals are 
shown in Fig. 44, a camera lucida drawing made while the canals 
were quickly disappearing from view. Three of Fraipont's large 
"canaux descendants" can be seen together with much of the anas- 
tomes among them and at least two branches to foramina secundaria. 
The largest and rno3t median canal was found to have a diameter of 
50^ . But contrary to vhat was stated by Fraipont (1881 : 3, 11) 
only 3ix of these main channels were found close to the ventral 
layer of longitudinal muscles in the meduallary parenchyma and not 
six for each face, Loennberg stated, correctly it seem3 to the 
writer, that their number is very variable as i3 their 3ize and 
course, the whole forming a complicated reticulum showing the 
typical "island formation". A3 regards the termination of the ex- 
cretory vessels at the posterior end of the strobila the writer was 
able to confirm Fraipont' s 1831 : 10-11) statement that: "Chez un 
3ujet qui a de'ja perdu des proglottis, les gros canaux longitudi- 
naux sont rompus au niveau du bord libre poste'rieur du dernier 
segment. Les uns communiquent directement avec l'exterieur, les 
autres ne sont renferrne's et 3e terminent en cul-de-3ac; " but no 
cases were met with in the material at hand in which it could be 
considered that no segments had been lost. Towards the 3colex the 
six ve33el3 gradually come closer and closer together until in the 
first segments they may appear for short distances in two sets of 



134 



three each, doraoventrally situated, lout soon again become lo3t 
in their anaatomasea. Entering the scolex four, three, or two 
main canals may be seen, but here they cannot be followed aa 3uch 
thru many aectiona since they aoon break up into the recitulura 
mentioned by Fraipont aa ramifying thruout the 3colex. 

Of the generative organs the earlieat writers were able 
to discern only the external openings ("oscula" and the uteri 
Which, showing their contained dark brown eggs thru the body wall 
aa a longitudinal series of dark punctationa, gave origin to the 
apecific names of Zeder (1300) and Rudolph! (1302 and 1310). Con- 
cerning these characters Mueller (1738 : 5) wrote as follows: 
"Marginea corporis depress! intersectione 
articulorum crenati apparent; oacula in aaaterior- 
ibua articuli8 nulla adsunt, in posterioribua vera 
altera in pagina pori in macula albida nigrieantes, 
in altera papi lla alba aubelevata, punctaque 3eu 
globuli utrinque diapalati, qui ovula . Oscula aeu 
pori non aeriem rectam in corpore. Taeniae sed 
"nine et illinc divergentem constituunt, alt.irum in 
centra articuli, alterum in intersectione const! tu- 
um. articuli postioi reliquis latiore3 punctis 
utrinque di3persi3 medio autem coacervatia papillu- 
lamque exhibentibua repleti aunt; harum coacervatio 
oculo nudo punctum centri nigrum offert, armatus 
vero ovula seu globulos e membrana ovata pfcllucida 
punctuli3 nigricantibua impleta conatantea di3cernit". 
Rudolphi (1810 : 50) described them in these words: 



165 

"Singuli enim articuli in 3uperficie 
dor3ali nodulum orbi sularem, simplicem vel dupli- 
cem, 3ubelevatum, vel albidum vel fusee 303 atern 
ant nigresoentem exhibent; in superfioie autem ven- 
trali nodulus simplex vel duplex, pariter, sed 
minus, exatan3, qua3i perforatus videtur; corpor- 
eque pellucido nodulis illis linea cor.- oris media, 
plerumque tamen irregularis oritur, Noduli aperti 
ovaria si stunt j via que elliptiois mediocribus re- 
ferti sunt, haeo etiam 3aepe circa eosdem effusa 
sunt . " 

From these descriptions it is to be seen, incidentally, 
that while Mueller made correct observations concerning the rela- 
tions between the positions of the genital openings and the trans- 
verse furrows mentioned above, Rudolphi considered the ventral 
surface to be that on which the openings of the cirrus and vagina 
are situated and the dorsal that on which the uterus opens to the 
exterior. Van Beneden (1350) seems to have been the first writer 
to describe the anatomy, with, however, some errors of interpreta- 
tion as pointed out by Loennberg (1891e). After Loennberg's the 
best and practically the only description of the genitalia was 
given by M&tz (1392 : 105-103), Ariola (1900 : 394-7) and Luehe 
(1910 : 25) obviously copying in part at least from him. 

The earliest traces of the reproductive rudiments appear 
in toto mount 3 of this form about 35 to 40mm. from the tip of the 
scolex. From this region backwards they increase in 3ize, but 
so slowly that in large 3trobila3 there may be an intervening 



1S6 



stretch of at least 335mm, before the genital 3inus appears. 
Then the rudiments differentiate vary quickly, the first eggs 
appearing in the uterus-sac about 2mm. farther on in one toto 
mount made. In the largest 3trobila at hand (no. 137 above) the 
first genital sinuses were seen, when the vorfri was examined in 
alcohol, about 375m.ii. from the tip of the scolex and the first 
traces of eggs showing thru the ventral body wall about 30mm. 
farther on. 

Van Beneden (1350 : 168) seems to have ben the first to 
mention the relations between the external segments and the sets 
of reproductive organs. He said: "Dans chaque anneau, il y a 
deux on trois appareils males et femelles complets ; je pense que 
ces anneaux se divisent encore plus tard, de maniere "a n ! avoir 
plus qu' un appareil dans chaque animal," — (the "animal" is 
evidently a misprint for "anneau"), — and further in his footnote 
referring to the superscript after "complets": "J'ai vu des anneaux 
qui en contenaient jusqu'a Six." In his Fig. 4, PI. XXI, he 
showed four parts of the strobila containing evidently three or 
four sets of reproductive organs in each segment, with the latter 
subdividing so that two sets appeared in each subsegment in the 
fourth part of the figure. Leidy (vide supra) described the pos- 
terior segments of B. scorp ii as " ... quadrate; each «ith an 
appearance of three subdivisions, with the subsegrnents having a 
pair of generative ap-rtures, in the course of a longitudinally 
depressed, dark colored line, passing the length of the body." 
Linton (1890 : 733) referred to "the phenomenon which the posterior 
segments present of being welded together in groups of three or 



167 

four, an appearance which i3 quits characteristic of the posterior 
segments and which has been alluded to in various descriptions of 
the 3pecies," while further, in connection the apertures of the 
reproductive organs: "In the middle of the strobila there sometimes 
appear to be as many as four or more papillae to a single segment, 11 
and with reference to the specimens from Lap hop se tta macula ta : 
" ... toward the posterior end of the body ths adult segments are 
arranged in groups of from four to six simple segments, as if the 
latter were partially fused together, which is another characteris- 
tic of this species." From these statement and the further fact 
that the posterior proglottides have been described a3 trapezoidal 
(Stossich), quadrate (Rudolphi, Linton), subquadrate (Die sing), or 
at moat, broader than long (Rudolphi), — actually about twice as 
broad a3 long from Linton's (1890 : 732, 734) description, — it 
is evident that the groups of four sets of reproductive organs 
(much less frequently three, five or six) 3hown here in Figs. 43, 
45 and 46, and separated from each other by grooves which in alcoho- 
lic material appear to be complete, have been considered to consti- 
tute the ripe proglottides. But, as pointed out d3} Loennberg, 
the lesser transverse furrows are only "greater wrinkles or foldings 
of the surface" and do not cut in deep enough to cause th: parts 
immediately ahead to stand out distinctly like the posterior borders 
of the proglottides of othsr species, e.g., 0. cra33iceps . Such 
posterior borders, with their accompanying "complete" transverse 
furrows, do occur, however, but only at considerable intervals. 
One case is 3hown in Fig. 47, where it will be noticed there is no 
such distinct separation of the proglottia from the next one ahead. 



168 



So far 3.3 the writer is aware, this has been pointed out only by 
Luehe (1910 : 25) when he said: n ... in reifen Gliederstreoken 
liegen zwischen zwei vollig durchgehenden, aber auch nur wenig 
hervortretenden oberf 1'achlichen Querfurchen in der Kegel 16 sehr 
kurze Genitalsegmente , die ausserlich voneinander nur duroh 
"ackenbildungen des Seitenrandes ge3oheiden sind." In this con- 
nection it should also be noted that in his description of 
Fimbriaria fascio lari s (Pallas), a taenioid from various water 
birds, Wolfflugel (1900 : 94) remarked that it ia comparable to 
B. scorpii in that "Eine bis ins aussergewohnliche gestiegerte 
Anzahl von Geschlechtsapparaten in einer Proglottis 3ich f olge . " 

In a considerable length of one to to mount of this form 
there were found ahead of and including the region of differentia- 
tion of the reproductive rudiments the following consecutive number 
(from behind forwards) of genital segments between the most pro- 
nounced transverse furrows, that is in the primary segments in 
question: 57, 82, 101, 107, 90, 111, 116, — using as the criterion 
of a genital segment, especially ahead of the region of differen- 
tiation, the aggregation of nuclei in the median line which will 
go to form the central organs and ducts of the system. But there 
is much difficulty in making these counts on account of rudimentary 
or intercalated groups of nuclei which, judging from conditions to 
be seen in theregion of differentiation, may or may not form complete 
3ets of genitalia, and above all of the further subdivision of 
many of these rudiments, which otherwise proceeds in quite the same 
manner (Fig. 42) as that of the external segments in the anterior 
part of the strobila. Furthermore, there may often be seen either 



169 

in the anterior part of the region of differentiation or much 
farther ahead (Fig. 42) a lateral doubling of the developing 
genitalia. But since no case was met vith of two a^ta of repro- 
ductive organs in a ripe genital segment, it was concluded, es- 
pecially because of the great infrequency of this duplication, that 
one or the other rudiment eventually gets the upper hand and 
develops at the expense of the other. This is borne out by the 
fact that in half the oases one rudiment was much larger than the 
other. The above mentioned groups of rudiments were found to be 
divided and subdivided by less and less pronounced transversa 
furrows in the following manner: 

•67 82 101 

43 24 35 47 54 47 

24+19 10+14 14+31 18+13+16 16+14+12+12 20+12+15 

This continued until eventually the groups of four (or five, rerely 
six) sets of genitalia of the author* could be made out. But 
these were seen to be divided into two groups of two sets each, so 
that each lateral crsnulation corresponded to two (or three) of 
these, i.e., to the l-32nd division described above (Fig. 32). In 
ripe segments this arrangement may obt in or the segment may divide 
again peripherally, so that each crenulation then corresponds with 
one set of genitalia (Figs. 45 and 46). The latter figures show 
that "complete" transverse furrows are present between every 8 or 
9 (sometimes 7, or apparently even 3, 4, 5 or 8!) genital segments. 
However, other more relaxed strobilas in alcohol showed complete 
furrows only every 13 ot 17 sets of genital segments, these often 



being a group of 5 instead of the much more common group of four, 
"but in the same neighborhood of the Strobila just as complete 
grooves every 8, 9 or 10 sets. This shows that a grouping of the 
genital segments into lots of approximately 13, aa mentioned by 
Luehe (1910 : 25) is not at all regular and can scarcely be said 
to occur even "as a rule". 

The genital sinus is situated on a low papilla (Fig. 47) 
on the dorsal surface and in the median line from one half to two- 
thirds of the length of the genital segment from it3 anterior 
border, while the uterine opening on the ventral surface is located 
much farther forward even at the bottom of the groove corresponding 
to the indentation of the edge of the strobila, separating the 
crenulations mentioned above. The sinus itself is circular in 
outline and from 40 to 45 p. in diameter by 15 to SO^jl in depth. 
kt its bottom the cirrus and vagina open close together, the latter 
immediately behind the former, thru a .secondary sinus or ductus 
hsrmaphroditicus, the walls of which are often found protruding t 
thru the opening of the larger vestibule as if to form part of a 
functional cirrus (Fig. 47). 

The testes are arranged in two lateral fiels in the 
medullary parenchyma, as pointed out by Loennberg, and are contin- 
uous from segment to segment, altho they show 3ome tendency towards 
division interproglotHdally . The number was given by Matz (1893 : 
106) as about 76, with their size as 40, 8p- , but here it was 
found to be 30-60, while their 3ize was 35 to 70 y~ , 60 being the 
commonest measurement. The vas deferens, filled with sperms, forms 
a compact mass of coil3 about 0.18 x 0.10mm. in size, lying 



171 

irregularly to the right or left of the uterine duct or slightly 
"behind the sac and immediately alongside the cirrus- sac, as shown 
in Fig. 48. The ductus e jaculatoriua portion of the vas deferens 
within the cirrus-sac, that is, that part occupying the lowermost 
one-third of the latter, has a diameter of 4 to 6|a. . The middle 
stretch of the duct often expands to 13^, while the distal part, 
the cirrus proper, has a maximum length of 65 ^ with a width of 
14y- . Matz gave the dimensions of the organ (? the cirrus- sac) 
as 100 x 50 f/L . The cuticula lining the cirrus is pseudociliated 
on it3 inner (functionally outer) surface, somewhat as is that on 
the external surface of the worm. The cirrus-3ac i3 located at 
right angles to the dorsal surface (Fig. 47) and extends only a 
short distance into the medulla, as compared to other species. It 
is ovoid in shape, with the narrower end towards the cloaca, and 
115-130 ia- in length by 75-80 in diameter. Its wall is composed of 
an inner thick layer of circular muscle fibres and a very thin 
outer layer, the fibres of which are directed somewhat obliquely, 
the whole being 8^ in thickness. As pointed out by Loennberg and 
shown in Fig. 47, the organ is peculiar in that its wall is coated 
both externally and internally with a thick layer of nuclei which 
are doubtless mostly myoblastic in their nature. An aggregation of 
nuclei at the lower pole of the^ac, surrounding the vas deferens 
and continuous with the layer of nuclei on the outside of the 
pouch, seem to be too numerous to be considered as myoblastic 
nuclei only. They do net seem to be mentioned either by Loennberg 
or Matz. Their arrangement would indicate that they are possibly 
prostatic in their nature, the whole structure having the appearance 



172 



of a gland. Retractor mu3cle9 of the cirrus proper are 3carce. 
Thi3 fact taken in conjunction with the further fact that the 
■rail of the 3ac i3 quite thick and powerful and that Loennberg saw 
only a short thick cirru3 when protruded, would lend support to the 
view that the latter is quite small and not very important from^. 
functional standi.: oint. Concerning copulation in this species 
Loennberg said: "Es ist daher wahrscheinlich, dass die normale 
Befruchtung so vor sich geht, d>ss das Sperma in den Sinus genitalis 
hinausgepre3st wird, und davon entweder passiv durch die Kontrak- 
tion der Sinus genitalis oder aktiv durch eigene Bev/egung in die 
Vagina gelangt. So^wohl die eine als die andere Wei3e 3cheint 
recht moglich zu 3ein, well die Mundung der Vagina ganz neben der- 
jenigen des Penis gele^en i3t." 

The vagina opens into the ductus hermaphroditicus close 
behind the cirrus, and from that point passes close along the cirrus 
3ac to it3 lower end, and then turns back to pass over the ovarian 
isthmus and into the generative space. It3 diameter is 1 3 p. , while 
its wall is composed of a cuticula 5^ in thickness and a thin layer 
of circular muscles. It 3 cuticula i3 retained until the point of 
union with the oviduct is reached, where the lumen narrows down 
suddenly to one half the former diameter (Fig. 49). The ovary is 
somewhat irregularly "bi 3 cuit- shaped" (Fig. 43) situated clo3e to 
the po3t3rior border of the segment or protruding slightly into the 
segment behind. It is composed of short tubular lobules of varying 
size and has a width of 0.35mm. and a length of 0.15mm. In trans- 
sections it i3 3een to be "concave towards the surface bearing the 
genital openings" owing to the fact that the ventrally situated 



173 

isthmus is quite narrow and thick and consequently not well 
separated from the lobular rings which extend thruout the whole 
dorsoventral diameter of the medulla and also somer/hat enfold the 
former posteriorly in the median line. Ova from the isthmus are 
15 u. in diameter, while their nuclei and nucleoli average, respec- 
tively, 7 and 3 1*- . The oocapt is quite muscular, and 35^ in 
diameter. The oviduct proceeds dorsally for a short distance only 
before it is joined by the vagina at a vestibule into which the 
oviduct itself opens (Fig. 49) by a narrow slit much as in CI. 
crassiceps . The wall of the duct is composed of an epithelium, in 
which no cell-boundaries could be made out, but provided with cilia 
directed towards the uterus. The oviduct continues dorsally for 
a short distance with the same structure and diameter, namely 
18^ , to take un the vitelline duct dorsal to the anterior edge of 
the isthmus. The vitelline follicles are arranged in the cortical 
parenchyma in two lateral fields (Fig. 48) which are, however, 
slightly connected with each other dorsally andjvent rally in the 
median line by a few isolated follicles. No large follicle such as 
that described by Matz in the neighborhood of the ovary was seen in 
the material studied. The follicles vary somewhat in size, but 
average 35 to 55^ in diameter, are very closely crowded together, 
— so as to obscure in toto preparations the testes beneath them, — 
and continuous from proglottis to proglottis. The latter fact 
makes it difficult, if not somewhat unnecessary, to state the number 
for each genital segment, but, using Matz ! s method of multiplying 
the average number seen in transections by that seen in Sagittal 
sections (here the average of several segments was taken), the 



174 

number was found to vary from 350 to 540, or 440 on the average, 
lotz gave 490 as the number. Two main vitelline ducts proceed 
from opposite 3idea of the genital segment and unit in the ante*o- 
dorsal portion of the generative space to form a very short common 
duct which from the amount of yolk it usually contains may act as a 
vitelline reservoir, altho the same function is shared even to a 
larger degree by the much coiled and distended proximal portions 
of the separate ducts. A few cases were met with in which small 
ducts laden with yolk came from follicles clearly belonging to the 
genital segment following. This condition is, however, not sur- 
prising in view of the continuous arrangement of the follicles 
themselves. The diameter of the common duct at its point of union 
with the oviduct was found to be about 8^ . The very voluminous 
shell-gland is situated dorsal to the ovarian isthmus close behind 
the cirrus-sac with a depth of 85 and width of 115 . The 
uterine duct is quite capacious since it is composed of may coils 
extending thruout the whole depth of the medulla immediately ahead 
of the ovary. Proximally it is lined with a 3yncitial epithelium 
which distally becomes much attenuated. While it is usually 
situated in the median line it may alternate from right to left as 
a whole according as the va3 deferens does so on the opposite side 
of the proglottis, the uteru3-sac in such cases remaining in the 
median line. As above noted, the uterus-sacs were called "ovaries" 
by the early writers. They were seen thru the body wall to be 
filled with the characteristic dark brown eggs forming dark patches 
or punctations, hence the specific names bipuncta ta and punctata . 
In this species the uterus-sacs w$re described by Rudolphi, 



Lenckart (1819 : 41), et al . a3 arranged in a single row, in a 
double row, or alternating thruout the strobila. Here they were 
likewise found to alternate irregularly from side to side (Fig. 46) 
(e.g., r, 1., L, r„ r„ L, r., 1., 1„ r,, r., 1., etc.) or to be more 
meuialj y situated (1., m., 1., m., m., m., m., 1., m., r., r., m., 
1., 1., m., m., etc.) but never in two rows, excepting in a very 
few immature genital segments (Fig. 42), unless the alternating 
condition in much contracted strobilas is considered as such. 
iMle the sac has a diameter of about 0.18mm. when the first eggs 
appear in its lumen, it may reach a length of 0.35mm. and a trans- 
verse diameter of 0.22mm. or about cne- sixth of that of the pro- 
glottis. The combined uteru3-sac and uterine duct may in many 
oases occupy more than one- third of the width of the segment. The 
hindermost segments, in which the uterus- sacs may be gorged with 
eggs to a diameter of 0.65mm., separate from the chain evidently in 
pairs, the lines of division taking place at the furrows between 
the larger crenulaticns mentioned above. No detached proglottides 
were found, however, free in the intestine of the host, altho 
Olsson (1867 : 55) recorded having found such, while Weinland 
(1858 : S) said that, according to Eschricht, the species "which 
lives in the scftlpin of the Baltic ( Oottus scorpin.3 ) throws off its 
whole chain of joints every year, and then sends out a new one from 
the neck" Like that of the distal portion of the uterine duct the 
wall of the sac is composed of a much attenuated epithelium from 
the basement membrane of which the nuclei, separated by 'wide in- 
tervals, project into the iuman like bosses. The uterus-opening is 
situated ventrally in the middle of the uterus-sac, and with regard 



176 

to the external segmentation either in the middle of the larger 
(double) segment or in the groove separating it from the next ahead 
or behind. Circular in outline and 50p^ in diameter, it is sur- 
rounded by an area of radiating nuclei, thought by Loennberg to be 
possibly of the nature of a gland for the secretion of a material 
of use in the passage of the eggs to the exterior. The actual 
opening is formed by the rupture of a membrane guarding the outlet, 
7/hich has a thickness of from 15 to 30 ^ . (Cf. C. crassiceps ) . 

The fresh egg is ellipsoidal in shape, dark brown in color, 
and measures from 36 to 80^ in length by 43 to 45 in transverse 
diameter. The shell was observed to be about 9^ thick in living 
material and not provided with an operculum. No mature eggs show- 
ing the six hooks of the oncosphere were met with in fresh material 
in the field. 

Nothing was discovered regarding the life- hi story of this 
form, not even possible intermediate hosts in the way of food 
contents, for the stomachs and intestines of the few sea-ravens 
examined were all found to be empty. Linton (1890 : 732) gave a3 
the food of Lophopsetta maculata and Limanda f erru^i nea, from which 
he recorded Dibothrium punctatum Rud., "several species of Annelids, 
fragments of Squilla, and several specimens of a species of 
Margarita . " No specimens smaller than about 25mm. in length were 
obtained, so that the appearance of the youngest strobila was not 
observed. According to Udinsky's abstract, ?il«t (19.06 : 191), 
working on B. scorpii from Raja clavata of the Black Sea (the only 
case of the species having been found .in a selachian fish, so far 
as the writer is aware) established the fact "dass seine Larven in 



177 

den ve3chiedenen Fischen oder Tieren, welche von Rochen ( Raja 
clavata ) gefres-en werden, sich befanden." 

From the foregoing description it is to be Been that thifl 
form ia very closely related to the B. scorpii (Muei: er) of Europe, 
aitho in many respects it is 30 different a3 to almost warrant the 
erection of a new species to accommodate it. However, on account 
of the fact that several forms of the European species have been 
reported, namely, B. scorpii forma buba l idis and forma motellae by 
Loennberg (1889 : 32 and 1893 : 13) and those from Rhombus maximus 
and Cottus ggadricornis by Schneider (1902a : 14-15 and 1903 : 
75-76), it was considered that here in America we have the same 
species as has been found in Europe. And from a comparison of 
the measurements given above with those given by Leidy (1855 : 444) 
and Linton (1890 : 732, 734 and 1897 : 430), it seems that, little 
as we can rely on external measurements, they also point to definite 
differences of habit as this worm is found in different host 
species on this side of the Atlantic. 

In the $able given below under B. clavicep s the important 
diagnostic data of this form are placed alon^ide those of the Euro- 
pean species for the sake of comparison. 

The material studied consisted of lots T*os. 191, 19S, 197, 
198, 287, and 288 of the writer's collection from the intestine of 
Hemitripteru s americanus (Gmelin), No. 17.57 of the Coll. Univ. 111. 
from the same host, and No. 17.56 of the same collection from 
Myo xo cep halua ? aeneus . 



Species 2. Bothrioc ephalic clavicep s (Goeze,1782) 





(Figs. 50 - 


55.) 








1722 


Vermis mnl t i merabris 
anguillae 


Leeuwenhoek 


1722 




490 


1780 


Taenia an^uillae (part 


. ) Mueller 


1780 




208 


1782 


Taenia claviceps 


Goeze 


1782 




414 


1786 


Taenia claviceps 


Batsch 


1786 


: 


211 


1786 


Taenia an^uillae 


Batsch 


1786 


: 


233 


1790 


Taenia ansuillae 


Graelin 


1790 


: 


3078 


1790 


Taenia claviceps 


Schrank 


1790 


: 


46 


1800 


Rhvtelminthus aneuilla 


3 Zeder 


1800 




215 


1801 


Taenia ciaviceos 


Rudolphi 


1801 


• 


103 


1802 


Taenia anffuiilae 


Bosc 


1802 




307 


lOU<J 


RVivt i t* 1 3 vl rpTic 


Zeder 


1805 




2 93 


loin 


jj u i 1 1 lUi/C y)i la. J.uo tiavi" 

ceos 


Run o 1 ■oh i 


1810 




37 


1816 


Bothrioc. claviceps 


Lamarck 


1816 


: 


582 


1819 


Bothrioc. clavicens 


Rudolphi 


1819 


* 


136,472 


1819 


Bothrioc. claviceps 


Leuckart 


1819 


: 


49 


1824 


Bothrioc. claviceps 


Mtzsch 


1824 


; 


97 


1844 


Bothrioc. ciaviceos 


Bellingham 


1844 




251 


1846 


Bothrioc. ciaviceos 


Dujardin 


1845 




618 


1848 


Bothrioc. claviceps 


Siebold 


1848 


• 


147 


1850 


Dibothrium claviceps 


Diesing 


1850 




589 


1853 


Bothrioc. claviceps 


Baird 


1853 




89 


1854 


Dibothrium claviceps 


Diesing 


1854 


: 


578 


1859 


Dibothrium clatoiceps 


Polonio 


1859 


» 


225 


1859 


Dibothrium claviceps 


Molin 


1859 




8 


1863 


Dibothrium ciaviceos 


Diesing 


1863 




241 


1867 


Bothrioc. clavices© 


Olsson 


1867 




56 



179 



1865 


Bothrioc . 


claviceps 


Carus 


1885 


; 120 


1892 


Bothrioc . 


claviceos 


Mat z 


1892 


: 108 


18 93 


Bothrioc. 


claviceos 


Olsson 


L8 93 


: 16-17 


1896 


Bothrioc. 


claviceps 


Ariola 


1896 


: 280 


1899 


Bothrioc. 


claviceps 


Luehe 


1899 


: 43 


1900 


Bothrioc. 


Qlaviceps 


Ariola 


1900 


: 393 


1902 


Bothrioc. 


claviceps 


Fuhrmann 


1902 


: 441,^*47 


1910 


Bothrioc. 


claviceps 


Luehe 


1910 


; 25 



Specific diagnosis; With the characters of the genus. Large 
cestodes up tp 540mm. long by 2-3 wide. Scolex small, elongate, 
but usually found contracted to an almost spherical shape; 4 ©-1.5 
mm. long by 0.3-0.5 wide at the middle. Prominent terminal disc. 
First segments thick, short and crowded; middle , oblong; posterior, 
or ripe proglottides , usually 2mm. broad by 0.5-0,7 long, often qua- 
drate, arranged in groups of two, between which the transverse 
furrow is not prominent. Other transverse furrows well marked. 

Cuticula l-2<> thick. Calcareous bodies very scarce. Main 
longitudibal muscles not in bundles. 4-6 chief longitudinal ex- 
cretory vessels. 

No genital papilla; genital cloaca funnel-shaped, midway be- 
tween anterior and posterior borders of the progxottis. Vagina 
opens immediately behind the cirrus-sac; no separation between 
common cloaca and hermaphroditic duct. 

Testes large, subspherical, averaging 58(x long, 64|jiwide and 
60 deep; 50 to 60 for each proglottis. Coils of vas deferens 
loose, close behind uterus-sac, 0.35mm. wide by 0.07 long. Cirrus- 
pouch ellipsoidal, 127-145 deep oy 81-104 in diameter, thin-walled. 

Ovarycorapact, 0.45-0. 55mm. in width, 0.055 in length by 0.18 



180 

in depth; isthmus quite thiol, ventral. Oocapt 30^ 8n diameter. 
Vitelline^ollicles not separated into two fields on either surface, 
450 to 720 in number, 45, 80, and 85|iin length, width and depth, re- 
spectively; vitelline reservoir large, 175 x G5f-. Shell-gland pos- 
terodorsal, alternating irregularly from right to left opposite the 
vas deferens. Uterine duct quite voluminous , between ovary and ut~e- 
rus-sac. Uterus-sac transversely elongate, occupying one third or 
more of the transverse diameter of the proglottis, usually larger 
towards the side bearing the opening; openings form a zig-zag ven- 
tral row. 

Egg, 58 - 33 long by 37-40 wide, without opercula; light in 
color, show only faintly thru the body-wall. 

Habitat: In the pyloric portion of the intestine of the hodt. 



Host Locality Collector Authority 

Anguilla vulgaris — Leeuwenhoek Diesing 1850 :590 

" 9 Zedep » » » 

H " Rennes Dujardin Dujardin 1845:618 

n " Patavia Molin Diesing 1863:241 

tt tr 0negasee,Huss- Kessler Schneider 1902:17 
ian Finland 

" " Sinus Codani Olsson Olsson 1 1867:56 



L.Halland, Swe- 
den 



n 



Venice Ninni Stossich 1891:8 

0stsee,Warne- Braun Braun 1891:55 

miinde and Unter- 

warnow-Rostock 



Anguilla vulgaris 



? " 
9 » 



" acut irostris 

n tt 

Anguilla anguilla 
n migratoria 

Iluraena anguilla 



cassmi 



Anguilla rostrata 



Eupomotis g ihbosus 

Gasterosteus bispi - 
nosus 



Genova 
Nancy 



Parona 
Prenant 



Konigsberg & Muehling 
Memel 

Peninsula of Schneider 
Porkala, Finland 

Lake Garda, Italy Largaiolli 

Ireland Bellingham 



Germany 



Gryphswald 



Siebold,ColL 
Erit.Mus. 



Kroyer 

Eorke 

Rudolph i 

Kais.-konij 
nat* lkab 



181 

Matz 1892:109 
Ariola 1900:394 
Zschokke 1896:818 
Muehling 1898:35 

Schneider 1902:15 

Ariola 1900:394 
Bellinghaml844:251 
Baird 1853:90 



Luehe 



1910:25 



HJorja, Scania, Olsson 
Sweden 

Lakes Halen & 
Refunds j on, Jemt- 
land, Sweden 

E8nan,Gulfl of " 
Bothnia 



Stiles & 
Hassall 1912:124 

Goeze 1782:414 

Rudolphi 1810:38 

Leuckart 1819:49 

Olsson 1893:16 



Naples 
Chamcook L 



Rudolphi 



, Cooper 
New Brunswick 



Walnut L.,Mich. H.B.Ward 
Woods Hole, Mass. V.N.Edwards 



Rudolphi 1819:472 
Cooper 

(the present paper$ 



132 



Since the earlier writers dealt merely with the external 
features of the species, their descriptions are of little com- 
parative value when the finer distinctions are at stake. In late 
years Ariola (18S6 : 230; 1900 : 393), Luehe (1899 : 43), Braun 
(1900 : 1676) and Fuhrmann (1902 : 44-1, 447) dealt with it from a 
systematic standpoint, and finally Luehe (1910 : 25) gave a short 
diagnosis, in placing it in his latest classification of the group; 
but all referred back to the original and evidently only adequate 
description of the internally anatomy, namely, that by Matz (1892 : 
108-110). Kere will be given only the most important specific 
data of value for a comparison of the form studied with the de- 
scription by the latter writer, since the accompanying figures 
illustrate many of the details sufficiently. 

In the material from Fupomotis gibbosus all of the anterior 
proglottides were found to be much broader than long, on account 
of the contraction of the strobilas, while those in detached pieces 
were from four to five times as broad as long, as shown in Fig. 54. 
Apart from Matz, Olsson (1893 : 16) and Luehe (1910 : 25) have 
noted secondary division of segments, while Dujardin (1845) ori- 
ginally said that "On remarque en outre que souvent les articles 
sont tellement unis deux a deux, que chaque couple parait n'enjfaire 
qu'un seul avec une ride transverse et deux apparsils genitaux 
I'unktevant l 1 autre." This pairing of the ripe proglottides 
(Fig. 54) is due to the manner of segmentation which was found to 
be like that described for B. scorpii , only quite regular since 
the reproductive rudiments appear relatively farther forward in 
the strobila and seem to be more stable in development. Concerning 



183 

the arrangement of segments for this species Luehe (1910 : 25) 
said that, "Zwei auf einanderf olgende Genitalsegmente ausserlich 
haufig nur vollkommen geschieden, indeasen fehlen durchgehende 
Querfurchen auf den Flachen nie auf so weite strecken wie bei 
B. punctatus . " On account of the great degree of contraction of 
the 3trobila at hand, provided with scolices, the primary segments 
were not followed with entire satisfaction very far beyond the 
scolex, but the first two were seen to be divided into four sub- 
segments each, — the first one, shown in Fig. 50, including the 
four segments to the (*) at the side of the figure, — with some 
indication of the next division which would result in eight seg- 
ments to the primary segment; the third into eight, and so on. 
There were indications posteriorly, however, that the primary 
segment consists of at least 32 genital segments or proglottides, 
but as in B. scorpii the furrows separating sets of 16, 8, and 4 
genitalia become almost as prominent as those between the groups 
of 32, while even those separating pairs are not as faint as Olsson 
(1893 : 16) stated and showed in his Fig. 1, Tab. II. At all 
events it should be emphasized that the furrows are more distinct 
and consequently the proglottides better defined, at least external- 
ly, than in B. scorpii . 

The following table gives the measurements of three of the 
largest specimens studied: 



184 



Number 


17.35.1 


17.33.2 


17.54.1 


Length of scoiex 


. 44mm . 


. 46mm . 


. 46mm . 


Breadth of terminal disc 


0.23 


0.20 


0.22 


1 at middle 


0.28 


0.30 


0.33 


Depth of terminal disc 


0.20 


0.20 


0.20 


" at middle 


0.26 


0.20 


0.40 


" posteriorly 


0.27 


0.27 


0.46 


Length of strobila 


155 


150 


43 


Maximum breadth 


2.9 


2.9 


2.0 


Width of ripe segments 


2.0 


2.0 


1.6 


Length of " " 


4.0-5.0 


4.0-5.0 


0.4-0.6 



Up to the time when Die sing (1863 : 241) incorrectly 
described the genital apertures as marginal and alternating, the 
only references to the reproductive organs of this species were 
to the uterus-sacs which were visible a3 faint punctations in the 
median line. Carus (1885 : 120) failed to correct Diesing's error 
so that it remained for Mfttz (1892 : 109) to give the first de- 
scription of the genitalia, upon which we can rely, in which, 
however, only the differences between them and those of B. scorpii 
were emphasized. The earliest traces of the reproductive rudi- 
ments were seen by the writer a'oout 5mm. from the tip of the 
scolex while the first eggs in the uterus-sacs came at about 55mm. 
While the opening of the uterus i3 wel] towards the anterior 
edge of the segment, that of the genital cloaca is midway between 
the anterior and posterior borders. There is no papilla, the 
opening being a low funnel-shaped depression in which there is no 



135 

division into an external common cloaca and an hermaphroditic duct 
(Fig. 55). The important data concerning the rest of the reproduc- 
tive organs are contained in the following taole, where only those 
of comparative value are given: 



136 



B. scorpii B. claviceps 
European data Data by writ er European data By write r 



Length 35-G00ram, 

Breadth 1-7 

length of scolex 0.9-3.0 

Breadth " " 0.3-1.7 

Breadth of pos- 4.0 
terior segments 



Length of same 



0, 



Number of genital 16 
segments per ex- 
ternal segment 



G77mm. 

3.35 

1.2 

0.35 

1.8 

0.35-0.35 
8 or 16 



90- 54 0mm. 

2-3 
0.5-1.5 

0.5 

2 

0.5-0.75 



155mm. 

2.9 

0.46 

0.30 

2.0 

0.5 



Less than in See 
B.scorpii text 



Number of longi- 
tudinal excret- 
ory vessels 

Number of testes 

Diameter of same 

Dimensions of 
cirrus-sac 

Number of vitel- 
line follicles 

Size df same 



Arrangement of 
same 



6,8,12 
76 

40.8 

100 x50 

490 
30-40 



30-60 
35-70 
120 x 80 

350-540 

35-55 



56 
36-47 
109 x 64 

462 



50-60 
60-70 
145 x 10,9 

450-720 



In 2 separate Dorsal filJds Dorsal Dorsal u- 
dorsal fields; slightly uni- fields u- nited;ven- 



Dimensions of 
eggs 



Arrangement of 



2 ventral 
fields weakly 
united 



50-80 x 40 



ted; 2 ventral nited; 
fields weakly ventral 
united fields 
weakly 
united 

66-80 x 43-45 50-60 



trals uni- 
ted to same 
decree 



58-63 x 
37-40 



uteri 



1 row, alter- 1 row, alter- 
nating, or 2 rows nating,or 2 rows 



187 



Diameter uterus Only small por- 

: diam. segment tion of diameter 

Longitudinal "Close together" 
muscles 



1 : 6 1:3 - I:?. 1 : |5 

In bundles Not in bun- Not in 
dies bundles 



138 



From the above comparison it will be seen that alt ho the 
individuals from Fupomo tis gibbosus (tho3e from which the data were 
taken) do not exactly agree with the European species, they are 
sufficiently close to justify their being considered the same. And 
this was made more certain to the writer by the examination of some 
fragments of the European form, obtained by Professor Ward from 
Dr. 0. Fuhrmann of Neuchatel, Switzerland, who took them from 
Angui 1 j a vulgaris in "North Germany". But it should be stated 
that in the latter material the cirrus-sac and ovary are smaller 
and the uterus- sac much larger, occupying more than half the dia- 
meter of the proglottis in many places; or the reproductive organs 
seem to become mature relatively earlier, differences in degree of 
contraction and relaxation being taken into consideration. 

The material studied consisted of No. 389 of the writer* s 
collection from An?ui l la rostrata , Nos. 17.33 and 16.456 from 
coll. Univ. 111., the former from E-upomotis gibbosua and the latter 
from Anguilla vulgaris (North Germany), and No. 17.54 of the same 
collection from Gastero3teus bicepinosu s . 

Species 3. Bothriocephalus Cdspidatus spec.nov. 
(Figs. 56 - 53) 

Specific diagnosis: With the characters of the genus. 
Medium sized cestodes up to 180mm. in length by 3.75 in breadth. 
Scolex large with very prominent terminal disc deeply notched 
surficially; bothria long and narrow and quite deep posteriorly 
giving the scolex when viewed laterally the appearance of an arrow- 
head; 3.3mm. long, 1.0 wide at middle 3.5 deep posteriorly. First 



189 

segments subcuneate and circular in transection, with prominent 
posterior borders; middle gradually "broaden until much wider than 
long; posterior two to four and a half times wider than long or 
1-2. 7mm. in width by 0.8 in length. Posterior end of strobila 
laauajly rounded, even when segments have already become detached. 

Cuticula 3.5 jju thick, subcuticula 5S<jl . No calcareous 
bodies. Longitudinal muscles not in bundles. Four main longitu- 
dinal excretory vessels. 

Genital cloaca median, halfway between anterior and posterior 
borders of proglottis, deep and funnel-shaped. Vaginal opening 
close behind that of cirrus; hermphroditic duct-obscure. 

Testes on each side separated into two fields by nerve 
strand, inner much narrowed than outer; 50 to SO in each proglottis; 
110,60 and 80jjl in maximum width, length and depth, respectively. 
Vas deferens a large compact mass of coils, elongate and laterial 
to cirrus-pouch, 0.23mm. long by 0.16 in width, alternates irre- 
gularly from right to left. Cirrus-sac very large and thin-walled, 
0.25 mm. in length (depth) by about 0.20 in diameter. Cirrus pro- 
truded, 135^ long by 85 in diameter. 

Ovary compact, with limbs often turned forward, 0.60mm. 
fide, 0.10 long and 0.13 thick; isthmus thick. Oocapt 25-30 in 
diameter. Vitelline follicles 800 to 1000; 70,50 and 45 ^ in 
maximum depth, width and length, respectively; occupying almost the 
whole of the cortex, strongly united lor sally and vent rally. 
Common vitelline duct long and narrow. Uterine duct confined to one 
side of the median line, opposite the cirrus-3ac, alternating irre- 
gularly from side to side. Uterus-sac spherical, occupying one- 



[ 



130 



third of the diameter of the proglottis; opening me iian, close to 
the anterior edge of the latter. 

Eggs ellipsoidal 62-65 )ll long by 42-45 wide, oncospheres 
not developed within uteri. 

Habitat: Caeca and intestine of the host. 



Ko^t 

Stisostedion vi treum 
( ty] s host) 



ooality 

Flat Rock L . , 
Muskoka Dt . , Ont . 

Giant's Tomb Id., 
Georgian Bay 



Collector \ ri ty 

Cooper Cooper 

(the present 



Stizostedion canadense 



Eiodon tergisus 



aj.030ides 



Sandusky, Ohio. H.J .VanCleave 

New Baltimore, Mich. H.B.Ward 
Port Clinton, Ohio " 
Put-in Bay, Ohio " 
New Baltimore, Mich. ■ 
Kansas City, Mo. H.M.Benedict 
Havana, 111. K.J. VanCleave 

Keokuk, Iowa H.B.Ward 



Type specimen : No. 174.2 of the writer's collection. 

Co- type : No. 174.3 of the 3ame collection, deposited 
in the Collection of the University of Illinois under the direction 
of Professor Henry B. Ward. 

Type locality : Georgian Bay, Lake Huron, off Giant's Tomb 

Island . 



191 

So far a3 the writer has been able to ascertain a descrip- 
tion of this apecies has not yet been published, nor have any 
bothriaeephalid oestodes been reported from the hosts listed above. 
The specific name cuspidatus , here chosen, has reference to the 
peculiar shape of the scolex as seen from the side : cuspis , an 
arrow-head . 

The first segments of this species show subdivision 
according to the same plan as that described for B. scor pii . Each 
primary segment was seen to be divided into two segments of the 
second order (Fig. 56) and farther back these again into segments 
of the third order, and so on, until, when the rudiments of the re- 
productive organs appear, the primary segment, whose boundaries can 
be recognized by carefully following along backwards from the 
3C0lex, contains 32 of them. This plan can be followed .veil into 
the region of dif f erantiation. There is not nearly 30 much irre- 
gularity, introduced by intercalated segments and the subdivision 
of others, as in B. scorpii , altho the same sort of dominance of the 
anterior ends of the major divisions over their posterior ends i3 
seen not only in the size of the subdivisions and that of the re- 
productive rudiments but, in the beginning of the region of differ- 
entiation, in the rate of differentiation of the common rudiment 
into the different proximal organs of reproduction. The latter is 
indicated in good toto preparations as well as in sections. As 
3oon, however, as the lumina of the uterus sacs appear the plan 
becomes obscured by the gradual enlargement of the posterior borders 
of the subsegments even to those of the fifth order; so that in 
turn we can 3ee defined, as we . follow them posteriorly, groups of 



192 

33, 16, 8, 4, 2 sets of genitalia (Fig. 30). And eventually t 
the posterior end of medium sized strobilas and for considerable 
^tre tones of the largest, these x^&irs become separated, and the 
segment contains only one set of genitalia. 

No statements can be made by the writer concerning the 
intermediate stages of the life history of this species. It may 
be 3aid, however, that many of the earliest formed segments are 
lost long before they become sexually mature, 3ince most of the 
youngest strobila3 are found lacking the end proglottis. Constric- 
tions at about the middle were present in many of them, a3 if the 
length of segments behind that region might be thrown off a3 a 
whole; but, since this was not a constant feature, it was considered 
to be due rather to the fixation of a wave of contraction passing 
over the strobila, such as may be seen in living individuals as 
well as in plerocercoids of other species of cestodes, e.g. Scolex 
polymorphus . 

The material studied consisted of 24 lots from the collec- 
tions of the University of Illinois (Professor H. B. Ward), of 
Dr.H. J. Van Cleave and of the writer, from the hosts as above 
listed. 



193 



Speci e 


s 4. Bothriocephalic manubrif ormia 


{ Linton , 


loo3 ; 




(Figs. 64-70) 








1889 


Dibothrium manubri forme 


Linton 


1389 


: 45S 


1890 


Dibothrium manubri forme 


« 


1890 


: 738 


1897 


Dibot fa r i um manub rl forme 


it 


1397 


: 429 


1900 


Bothriocephalus manubrif ormi 3 


Ariola 


1900 


: 410 


1903 


Bothriocephalus manubrif ormi 3 


Porona 


1903 


: 7 



Specific diagnosis: With the characters of the genus. 
Large ceatodes up to 330mm. in length by 5mm. in maximum breadth. 
Scolex large, elongate, with prominent terminal disc deeply notched 
laterally as well aa surficially, constricted posteriorly; length 
3- 3. 5mm., depth at middle, 1.0, breadth of disc, 1.0. Bothria 
long and very narrow posteriorly where the walla are quite thick. 
Firat aegments cuneate with salient posterior borders which are 
distinctly emarginate; middle, broadly cuneate, les3 emarginate; 
poaterior or mature many time3 broader than long and closely crowded 
5 x 0.3mm.; gravid proglottides, 2 x 0.4mm. Posterior half to 
t-o- thirds of the strooila provided with a median line (the com- 
bined uterus-sacs) . 

Cuticula 4.5f^ thick. Calcareous bodies large, 18-36 x 
ll-15p. . Longitudinal muscles well developed, in bundles. An- 
teriorly 4 chief excretory vessels. 

Genital cloaca median or slightly displaced towards either 
side, deep and narrow, separated from the hermaphroditic duct by 
a narrow muscular velum, half way between anterior and posterior 



194 

borders of the proglottis. Vagina opens immediately behind cirrus 
or very slightly to one side. 

Testes ellipsoidal in shape, 64-75 ^ wide, 45-60 long, 
S4-30 deep; 50 to 70 in number, dorsal in the medulla. Vas deferens 
closely applied to inner end of cirrus pouch, 85^ long, 175 wide 
and 400 thick, somewhat crescentric in the dor30ventral-tran3ver3e 
plane, opposite the uterus-sac. Cirrus-3ac long and cylindrical, 
0.50 x 0.14mm., inner half deflected towards the vas deferens, 
walls very thick, com-osed mostly of circular muscles. Cirrus 
short, usually not extending out3ide of the proglottis, 30 to 35 (l 
in diameter. 

Vagina with bulbous sphincter near its opening, 50 (a. long 
by 70 in diameter. Ovary irregularly brnnched but compressed 
anteroposterior^ , 0.45mm. wide; isthmus only ventral. Oocapt 
30^ in diameter. Vitelline follicl s extremely numerous, 35 
long, 60 wide and 85 thick. Vitelline reservoir large, 60 ^ in 
diameter. Uterine duct voluminous on both sides of the median line, 
crowding all other organs. Uterus-sacs alternate irregularly 
from side to side, each 0.45mm. in diameter, encroach greatly on 
neighboring segments, with\fchick mu3CUlo-glan:lular fvnnel- shaped 
ventral portion. Apertures form two lines on the ventral surface 
1mm. apart. 

Eggs 58 x 34 ^, dark brown showing thru wall3 of uterus- 
sacs . 

Habitat : Inte3tine of the nost. 



135 



Ho3t Locality Collector Authority 

Tetrapterus albidus Linton Linton, 1889 : 458 

(type host) - ~ Woods Hole, Mass. 

Hi3tio phoru3 gladius Newport, R.I. " ■ 1830 : 731 

Istiophorus nigricans Woods Hole " " 1901:448 



Tetrapterus imper ator " ■ " " : 447 

( = T albidus)" 

Tetrapterus fie lone Portoferrajo,Id. Damiani Parona 1302 : 7 

Elba 

Type specimen : No. 4711, Coll. U.S.Nat .Mus. 

Co- type : No. 16.461, Coll. Univ. 111. 

Type locality : "?eneke3e?" 



As regards the segments of this species Linton (188S : 
456) stated that, "Immediately back of the head the segments are 
very narrow, and for a greater or less distance, depending on the 
state of contraction, maintain about the same width as the base 
of the head. In some individuals the small anterior segments con- 
tinue much farther back from the head than in the one figured. The 
segments are alternately short and long. This characteristic is 
quite plainly marked in those segments which immediately follow 
the head, is still noticeable on the median segments and also on 
the posterior one 3, but is not so plainly marked on the latter as 
on the two former." This is due to the plan of subdivision of the 
segments, which is quite like that described for the preceeding 
species. It can be followed with certainty, however, only in 
the "anterior" and "median" portions of the strobila and not pos- 
teriorly where the segments are very short and crowded close toge- 



196 



ther longitudinally even tho the latter may not show the rudiments 
of the reproductive organs. Fig. 66 is of a primary segment (the 
fifth from the scolex in this case) to 3how this method of segmen- 
tation. Here there is a marked dominance of the anterior over the 
posterior half of the segment as regards the rate of division; and 
this is seen to be also quite applicable to the subsegments even 
to those of the fourth order. "In one specimen examined," to con- 
tinue quoting from Linton, "the first six segments did not show 
this alternation in size. In the next fourteen segments, however, 
the alternation was quite evident." This indicates that he was 
well acquainted with the division of the segments into subsegments, 
but did not hit upon thejpxact manner in which it is carried out. 

A3 regards internal segmentation the writer found that, a- 
part from the sets of genitalia, the musculature in general, — 
and for that matter the arrangement of the vitelline and the testes, 
— bear out Linton' 3 (1SS0 : 731) conclusion that, "So far as any 
internal characters go, the body is practically continuous." 

Nothing is known of the life- history of this species. 

The material studied consisted of two lots: No. 4711, U.S. 
N.M., from the rectum of Tetrapterus sp. from "Penekese?" deter- 
mined by Linton, and No. 16.461, Coll. Univ. 111. (H. B. Ward) from 
the intestine of Histiopho rus gladius, obtained from Professor 
Linton and evidently the actual specimen described by him in 18S6. 
The species 7/as determined and the above diagnosis prepared from 
confirmatory sections of the latter. 



197 

Bibothr ium lac inia tum Linton 

Linton (1897 : 435) established this species on the basis 
of the material contained in lot No. 4741 of the collection of the 
United States National Museum from Tarpon atl anticus, the Tarpum, 
and again reported it from the same host species in 1901 (p. 437). 
Luehe (1899 : 43) in his list of the species of the genus 
Bofrn riocephalus s.str. remarked that "Von weniger gut bekannten 
Arten gehoren anscheinend noch nisrher Both riocephalus lacinistu3 
(Lint.) und occidentalis (Lint.)"; while Ariola (1900 : 414) also 
placed it in the same genus as he conceived it to be constituted. 

During the study of B. manu brif ormi a the writer was im- 
pressed with the great resemblance between D. laci niatum and it, in 
all but a few details the two being, in^fact, identical. The mea- 
surements for length and maximum breadth, as shown in the compara- 
tive table below, agree, while those of the scolsx and anterior 
segments are as near as can be expected from cestode material -.vhich 
is found in various degrees of contraction and relaxation. All of 
the conditions represented in Linton's (1897) Figs. 7-13, PI. XXX, 
were observed in the material of B. manubrif or mis studied, — when 
such obvious errors as, "Fossettes marginal as to head, correspond- 
ing to the flat surface of body," are taken into consideration, — 
while the description of the external features, excepting that of 
the posterior segments, applied injdetail. But later lot No. 4741, 
U.S.N.M., was obtained by Professor Ward, and the writer learned 
that his suspicions were well founded; for D. lacinia tum proved to 
be identical with B. manu b rif or mi s . The posterior segments "with 



198 

breadth one and a half times the length" had different proper ticna 
from those observed in mature material of the latter species, be- 
cause they were, altho gravid, of quite younger strobilas. The 
material of No. 4741 is, in fact, intermediate between No. 4741 of 
B. manu briformis and the 16.461 of the same species studied by the 
writer, not so much in size, since it does not show the regions 
so well, a3 in degree of maturity. The fact that "the segments 
are not uniform; one segment with a salient posterior border 
followed by about two With less salient borders is due to the irre- 
gular manner in which the primary segment divides into subsegments 
(vide supra). The dimensions of theeggs corres_ond, while the 
measurements of the cirrus-bulb, vaginal sphincter, and calcareous 
bodies are the same in the two 3pecies: Linton 3tated that in 
D. laci niatum "The reproductive cloacae lie along the median line 
of one of the flat surfaces of the body. The external openings of 
the uterus lie along the median line of the opposite surface." 
While the former was found to be the case, the latter was not, for 
the openings of the uteri lie irregularly on either side of the 
median line as in B. man ubri formi 3 ♦ Furthermore the cirrus-bulb 
was not found to h-^ve "its inner end deflected to the right (left, 
7vhen we take into consideration the fact that the common genital 
cloaca of D. laciniatum was considered to open on the ventral in- 
stead of the dorsal suface) where it communicates with the vas de- 
ferens, which lies in numerous folds in front and to the right of 
the cirrus-bulb," but to alternate irregularly from side to 3ide 
according as the uterus- sac and distal end of the uterine duct 
occupy the other side of the proglottis, while the vas deferens is 



199 

as given above. Altho the vaginal bulb was found to be a little 
larger in the material of D. lacini atum, its structure and position 
were also quite as in B. manubrif ormis . On the other hand no 
muscle fibres completely encircling both genital apertures, 3uch as 
shown in Linton's Fig. 5, PI. XXXI, were 3een, but what might easily 
be taken for such were formed by the crossing of much curved and 
spread longitudinal and transverse fibres of the body wall, in such 
a manner that the portions intersecting at the four corners run in 
almost circular directions and concentrically parallel to each 
other so as to give the appearance of the whole forming a complete 
ring in each case. The genital cloaca was found to be shallower 
than in the material from Hlstrophorus gla dius , which is evidently 
due to the fact that the proglottides were younger and not yet 
gravid as in those from the latter host. The uterus-opening was 
not found to be "lined with cilia" but with irregular rugged pro- 
cesses which are evidently only portions of the lining of the 
developing funnel and the external duct of the same. Finally the 
position and structure of the ovary, of the vitelline reservoir 
and of the various layers of the body exactly correspond in the two 
forms . 

Consequently the writer feels that there can be no doubt 
whatever concerning the identity of D. lacin iatum with B. manubri- 
f ormis , which fact would also seem to be recognized in the Fauna 
of the Woods Hole Region (Sumner, Osborn and Cole, 1913 : 5S5) 
where the :ornier is not found amon^the ce stories, altho the host, 
Tarpon atlanticus_, is listed, and since B. manubrif or mi s was de- 
scribed before D . lacinia tum, the latter must be considered as a 
species delenda. 



300 

®« histiophorua Shipley 

The writer would also like to call attention in this place 
to the fact that Shipley's (1901) Bothriocephalic histiophorus 
agrees in all essentials with B, manub rif ormis , which is almost to 
be expected since both are found in the same h03t genu.3. 

The description anci figure of the scolex is that of the 
latter species, altho the true nature of the bothria was not as- 
certained by Shipley on account of their almost closed condition, 
which was also seen in many specimens of B. manu b r i f o rm i s by the 
writer. Consequently it was described, obviously erroneously, 
as " ... provided with longitudinal 3lit-Iike depressions which 
hardly attain the dignity of suckers situated in the dorsal and 
ventral plane," The external features of the strobila are the 
same in both species, altho Shipley was describing a comparatively 
young specimen, as shown in his measurements of the scolex and in 
his figures showing the size of the uterus-sac. The description 
and figures of the genitalia, agree in almost all details. It i3 
quite apparent, ho*>vever, that his Fig. V, diagrammatic it is true, 
is entirely misleading as to the proximal connections of the repro- 
ductive ducts, one of which, the ootype, be confused with the isth- 
mus of the ovary. The ova in the latter were also found by the 
wri+er to be 15p- in diameter in B. manubrif ormis as in B. histio- 
phorus . Besides, his description of these central connections of 
the genital ducts is certainly not that of the genus Bo thr i o ceph alu3 ; 
for in dealing with the isthmus of the ovary, which he called the 
ootype, he said that "Into this region opens the small shell-gland, 



301 



and the ducts of the yolk glands. The shell-gland lies posteriorly 
to the ovary between the right and left halves of that organ and 
with the ducts of the yolk glands it opens into the ootype poster- 
iorly"! The measurements of the eggs and the description of the 
u x erus agree with tr.ose of Linton's species, excepting that the 
opening of the uterus-sac " ... does not seem to be provided with 
anything of the nature of a sphincter muscle .... " Altho the 
material at hand did not permit of the sectioning of such young 
stages in the development of the uterus-sac, it would seem from the 
somewhat varying nature of its funnel-shaped ventral end, described 
above for B. manu brif ormi s , that in more anterior proglottides 
it might be in such a condition as to be easily overlooked. Here 
Shipley makes a statement concerning the probable disposal of ripe 
eggs, which seems to the writer to be the natural conclusion to 
arrive at after a study of the varying contents of the uterus-sacs 
along the strobila, namely, "From what I have seen I think it pro- 
bable that eggs pass out from the tapeworm into the alimentary 
canal of the host and that in B. histiophorus the eggs pass freely 
out from each ripe proglottis and do not wait until the posterior 
proglottides break off to make their escape from the parent." The 
nature and arrangement of the vitelline glands, the vagina and its 
bulb or sphincter, the testes in number and position, and finally 
the cirrus-sac, all considered in connection with his Figs. I-IV, 
force the writer to the conclusion that, so far as can be determined 
in the absence of material for study, Shipley's B. histiophorus n.3p 
is identical with B. manubrif ormi s (Linton). 

In the following table a number of important measurements 



303 

of B. manubrl f orml 8 , D . laoinlatum and B. histlophorue are given 
for the sake of comparison. 



303 

D. lac inia turn B. histiophorus B.man uhrif ormhs 

Maximum length of 154 mm. - 220 mm. 

strobila 

Maximum breadth of 4 5 

strobila 

Breadth at posterior 2 2 

end 

Length of scolex ? 1.8 1.5-3.5 

Breadth of terminal 0/8 0.4 0.8-1.2 
disc 

Ereadth of scolex at 0.4 0.G4 

middle 

Breadth at posterior 0.6 0.81 

end 

Breadth at constriction 0.25 0.21-0.44 

Depth of terminal disc 0.5 0.89 

Depth of scolex, middle 0.55 0.90-1.05 

Depth at posterior end 0.35 — 0.63 

Depth at constriction 0.25 — 0.58 

Length first segment 0.7 0.39 

Breadth same anteriorly 0.3 0.28-0.54 

Breadth same posteriorly 0.65 — 0.50-0.89 

Length of median seg'ts 0.3 0.3 

Breadthof median segHs 0.3 

Length posterior seg'ts 1 0.16("ripe" ) 1.0 

Breadth of same 1.5 0.5 ("ripe") 2.50 

Length of cirrus-sac 0.4 0.50 

Max. diameter of same 0/14 0.14 



;04 



Length of vaginal sphin- 0.05 0.05 

cter 

Diameter of same 0.07 0.07 

Dimensions of eggs 52 x 35 45 x 35 58 x 34 

Dimensions of calcar- 17-24 x 18-26 x 

eous bodies 8-14 11-15 

Number of testes 50 - 79 60 - 70 

Diameter of ova in 0.15 0.15 

ovarian isthmus 



305 

Species 5. BothriooephaluB oooidentalla (Linton, 18S7) 
(Figs. 71 and 73) 

1897 Dibothrium ocoi dentale Linton 1897b : 437 

1899 Bothriocephalus oooi den talis Luehe 1899c : 43 

1900 Bothri oc epha lus Occident all a Ariola 1900b : 415 

Specific diagnosis: With the characters of the genus. 
Large cestodes with maximum length of at least 310mm. and breadth 
5,5. Scolex small, elongate and somewhat rectangular, constricted 
posteriorly, 1.3mm. long by 0.46 wide. First segments somewhat 
funnel-shaped; middle, densely crowded, ten to twenty times broader 
than long, posterior narrower and longer, 3 x 0.8mm., in groups of 
three or four. 

Cut: cula 1.5p in thickness. Calcareous bodies 18 x 13^ . 
Longitudinal muscles in bundles, outer series very scarce . Four 
chief excretory vessels, two much more prominent than the others. 

Genital cloacae form a nrrow zig-zag row, each very shallow, 
no velum, cloaca and hermaphroditic duct united. Vagina opens 
directly behind cirrus or a little to one side. 

Testes divided into two fiels on each side by the nerve 
strEtnd, 75 to 30 in number, 35, 85, and 115pu in average maximum 
length, breadth and depth. Coi'is of vas deferens loosely arranged, 
the duct 35 [*/ in diameter, alternating; irregularly from side to 
side opposite the uterus-sac. Cirrus-sac long and cylindrical, 
0.33 x 0.06mm., walls comparatively thin, most of circular muscles 
being towards the inner end. 



206 

No vaginal sphincter or bulb. Ovary solid, unbranched, 
0.5-0. 6mm. wide, 0.04 long and 0.13-0.18 deep. Oocapt 25i>u in 
diameter. Vitelline follicles very numerous, in two lateral fields 
on each surface, leaving a broad median strip free, 25, 60, and 
115 in length, breadth and depth, respectively. Vitelline reser- 
voir 45 |j~ in diameter. Uterine duct voluminous on both 3ides of 
the median line, crowding all other organs. Maximum width and 
length of uterus-sac, 0.65 and 0.35mm., respectively; not en- 
croaching much on neighboring proglottides; ventral portion not 
especially modified, Uterus-openings alternate irregularly from 
side to side near the median line, far forward in the proglottides. 

Eggs 72-76 x 38-41 ^ , dark brown, showing thru the walls 
of the gorged uterus-sacs. 

Habitat: Intestine and pyloric coeca of the "rock cod", 
Seba s todes sp? 

Type specimen : No. 4740, U.S.N.M., collected by T. H. Bsan 
and identified by Professor Edwin Linton. 

Type locality: Whatcomb, Washington. 

The material contained in No. 4740, U.S.N.M., u: on which 
Linton based the species, was examined by the writer and confirma- 
tory sections made of mature segments, but it was all in such a 
very poor state of preservation that much of the detail could not 
be made out. However, as the specific diagnosis indicates, this 
is a true species of the genus Bothri ocephalic s.str. and quite 
distinct from B. manubrif ormi s which it closely resembles in many 
respects, internally as well as externally. So far as the writer 



307 

Is aware it is the only bothriocephalic! that has been reported for 
the Pacific coast of America, the fauna of which in this connection 
h:;s thus only been touched. 



308 

Genus 2. Cles tobothrium Luehe, 1899. 



Bothriocephalic (part.) 


I udolphi , 


1819. 


Dibothrius (part.) 


Rudolph i , 


1819. 


Bothriocephalus (part. 5 


Leuckart, 


1819. 


Bothriocephalus (part.) 


Dujardin, 


1845. 


Mbothriua (part.) 


Diesing, 


1850, 


Dibothrium (part.) 


Molin, 


1858. 


Dibothrium (part.) 


Diesing, 


1863. 


Bothriocephalus (part.) 


Carus, 


}885. 


Bothriocephalus (part.) 


Aridla, 


1896. 


Clestobothrium 


Luehe , 


1899. 


Bothriocephalus (part.) 


Ariola, 


1900. 


Clestobothrium 


Braun, 


1900. 



Generic diagnosis: Scolex almost spherical, the free edges 
of the dorsoventrally situated bothria fused with each other in 
their whole extent, in such a manner that only a small surficial 
opening near the apex leads into the interior of the spacious, 
hollow organ of attachment, flattened in the sagittal direction, 
by means of a short almost sagittally coursing canal which can be 
closed by a sphincterwjike musculature. External segmentation 
complete. Vitelline follicles in the cortical parenchyma. Ovary 
median and ventral. Receptaculum seminis small. Beginning of the 
uterus a winding co.nal which leads into an extraordinarily spaci- 
ous uterus-sac, distorting all the other genital ofcgans in ripe 



309 

proglottides. Uterine opening about median as is the dorsal geni- 
tal opening. 

Type species: C. crassiceps (Rudolphi). 

Clestobothrium crassiceps (Rudolphi, 1819) 



1819 


Bothriocephalus crassiceps 


Rudolphi 


1819 : 


139,476 


1820 


Eothriocephalus pilula 


Leuckart 


1819 : 


45-46 


1845 


Bothriocephalus crassiceps 


Dujardin 


1845 : 


617 


1850 


Dibothrium crassiceps 


Diesing 


1850 i 


587 


1858 


Dibothrium crassiceps 


Molin 


1858 j 


134 


1863 


Dibothrium crassiceps 


Diesing 


1863 i 


2 36 


1885 


Bothriocephalus crassiceps 


Carus 


1885 ; 


120 


1896 


Bothriocephalus crassiceps 


Ariola 


1896 j 


280 


1899 


Clestobothrium crassiceps 


Luehe 


1899 


44 


1900 


Bothriocephalus crassiceps 


Ariola 


1900 


397 


1900 


Clestobothrium crassiceps 


Eraun 


1900 


1692 


1901 


Dibothrium crassiceps 


Linton 


1901 


411,451 
473 


1909 


Dibothrium crassiceps 


Johnstone 


1909 


87-89 



Specific diagnosis: ^ith the characters of the ^enus. Medium 
sized cestodes, up to 92mm. in length, with a maximum breadth of 
1.5mm. Anteriorly surface of body with closely arranged transverse 
furrows, posteriorly segmentation more distinct, serrate. Scolex 
globose, 0.64-1. 08mm. long, 0.52-0.90 broad, andO. 68-1. 21 thick; 
divided by longitudinal marginal grooves into two dorsoventral 



hemispheres, the bothria. Latter, large, prominent, oval, their 
apertures about one-third from the apex and connected by a sadcUe- 
shaped groove over the tig of the scolex, with prominent lips. No 
neck, segmantatiom beginning immediately behind the scolex. Young 
segments closely arranged, five to six times as broad as long; 
mature proglottides quadrate to twice as long as broad, frequently 
divided on one or both sides by spurious articulations usually 
behind the uterus-sacs. 

Cuticula 2 to 5f^ thick, subcuticula 20. Chalk-Bodies absent. 
Musculature well developed, powerful sphincter around orifice of 
bothrium. Chief nerve strands ventral, 15 to 20^ in diameter. 
Usually four longitudinal excretory vessels. 

Genital cloaca median, dorsal, three-fourths to one-half the 
length of the proglottis from its anterior end, usually just pos- 
terior to the spurious articulations; hermaphroditic duct within 
this. 

Testes in two lateral fields in the medulla; ellipsoidal in 
shape, 0.125mm. long by 0.04 in diameter, continuous from joint to 
joint, 40 to 50 to each proglottis. Vas deferens forms a wedge- 
shaped mass of coils ahead of cirrus-sac and alongside of the hin- 
der end of the uterus-sac. Cirrus-sac elliptical to somewhat ova}, 
0. 128-0. 162mm. long by 0.087-0.116 wide and 0.098-0.116 deep, im- 
mediately behind the uterus-sac or lateral to its poaterior end. 
Cirrus-sac and vas deferens together alternate irregularly from 
right to left opposite the hinder end of the uterus-sac. 



311 

Opening of vagina close behind that of cirrus. Receptaculum 
seminis present as a short diverticulum almost parallel to the ovi- 
duct at the point of union of the vagina with the latter, about 10^ 
in diameter. Ovary bilobed, the isthmus narrow and ventral, ova in 
same 18 x 10^. Oocapt 20^ in diameter. Vestibule at the point of 
union of the vagina with the oviduct. Vitelline duct expands into 
a reservoir 30^ in diameter. Vitelline follicles not in, lateral 
fields, but continuous from joint to joint, 60 x 30 x bOy- in dimen- 
sions, about 700 in each proglottis. Uterus-sac elliptical in out- 
line, directed anteroposteriorly in the naterior half of the pro- 
glottis where in gravid segments it occupies almost the whole ofl 
the medullary region; 2.20 x 1.34mm. in dimensions; in quadrate 
segments irregularly alternating from side to side as are the uter- 
ine openings. 

Eggs, 75 x 40 j^. 

Habitat: In tha anterior portion of the intestine df the 
host. 



Host 

Gadus merluccius 
[Type host) 



euxmus 



Merlangus carbon - 



Locality 

Naples 

Patavia 
Trieste 
Trieste 
Nizza 



Collector Authority 

Rudolphi Rudolphi 1819:139 

Leuckart Leuckart 1819:45 

Molin Molin 1858:134 

Stossich Carus 1885:120 

t? Stossich 1899:1 

Wagener "'rgener 1854:61 



Merlangus sp, 



r agener 



Merluccius bilinearis V. r oods Hole Linton 



esculentus 



Trieste 
merluccius Pisa 

Ireland 



vulgaris 

tr 



Pisa 
Padova 
Patavia 
Genova 



Parona 
Stossich 
Wagener 
Drummond 
Wagener 
Kolin D 
Molin 
Parona Ari 



Portaferr- Damiani 
ajo,Id.21ba 



Pisa 

Gaeta 

Augusta, 
Catania 



Parona 

Ariola 

Earbagallo & 
Drago 



Pomatomus saltatrix Woods Hole Linton 



'A small hake" 



Calf of Man, Johnstone 
England 



Merluccius bilinearis Passamaquoddy Cooper 

Bay, St. An- 



drews, H.B. 

Buzzards Bay, M 
Mass. 

Vineyard Sound " 
Mass. 

Casco Bay, Me. n 
South Harpsv/ell 



YTagener 

Linton 

Ariola 

Stossich 

Wagener 

Thompson 

Diesing 

Diesing 

Molin 

Ariola 

Parona 

Parona 
Ariola 



213 

1857:93 

1901:473 

1896:265 

1898:115 

1854:68 

1844:439 

1863:237 

1863:237 

1861:235 

1896:265 

1899:8 

1899:8 
1900:397 



Earbagallo 
Drago 1903:412 

Linton 1901:451 

Johnstone 

1909:87 

Cooper 

(the present paperi 



313 

In external appearance this species is characterized by the 
globose nature of the scolex and the serrate margins of the strobila, 
the former of which was the basis of Leuckart's (1819 : 45) speci- 
fic name and which with the latter was emphasized and included in 
the diagnoses of all the authors after Rudolphi (1819 ) . But 
another important character which also assists in the ready recog- 
nition of the species is the presence of spurious articulations, 
which, however, are evidently not those mentioned collectively by 
Wagener (1854 : 59) as "articulatio spuria" (vide infra). The 
scolex (Figs. 73-77), as noted above, is divided by two longitudinal 
marginal grooves into two dorsoventral hemispheres, the bothria. 
The latter were considered by Rudolphi (1819 : 139, 477) ana others 
to be marginal or lateral (but not "lateral") in position, which 
error was finally and definitely corrected by Luehe (1899 : 35); 
but Leuckart (1819 : 45-48) rightly described and figures the 
scolex as "medio marginali sulcato, foveis lateralibus ... " and 
"Die Randflache des Kopfes ist breiter als die Sei tenf lache , die 
Mttelfurche jener ziemlich tief, und bildet an jener Seite eine 
erhabene, in der Mitte hellere Wolbung." It seems that Molin 
(1881 : 235) fell into the ervor of considering the marginal or 
lateral grooves, separating the bothria, to be the bothria them- 
selves, as indicated in his diagnosis: "Caput magnum subglobosum, 
utrinque sulco longi tudinali laterali, apertura centr-?.li bilabiata 
antica, bothriis ovalibus, 3ubterrninalibus , marginalibus , long! s", 
and in his "Osservazione 2" he said: "Ouantunque la testa sia molta 
grossa ad opaca, cib non per tanto potei distinguere il 3olco 
menzioaato da Diesing (1850 : 587) il quale perb corrisponde at la- 



314 

ti e non ai margin! del corpo, e sembra dividere la testa in due 
emi3feri. Ognuno di questi porta una f03sett& oblunga, ovale, 
che si estende dall 1 apice a due terzi della lunghezza del corpo, 
e sembra di quattro quadranti suddivisi da due solchi che s'in- 
crocciano." It is evident from his Fig. 2, Taf. V, that the 
"fossetta oblunga" is the entrance to the bothrium, but he does 
not seem to have observed the actual opening, not even in either 
marginal sulcus! Matz (1892 : 103) expressed the opinion that 
the bothria of this species were dorsoventral in position, while 
Ariola (1896 : 280) evidently on the basis of former diagnoses, 
placed the species among those of the genus Bothricephalus Rud. 
with "Botridi marginali". Stossich (1838 : 115) also described 
the scolex as " ... 3ubglobosa, con botridii marginali, subter- 
minali , ovato-allungati and Ariola (1S00 : 398) finally correc- 
ted his own view of the external structure of the scolex by saying 
that "Un esame anche superficiale dimostra pero che la posizione 
degli or?:ani di fissazione non e quale fu ritenuta, per che ciascun 
d'essi corrisy onde ad una faccia larga dello strobila, o como si 
dice, 3ono dorsoven^r.-.li . I pretesi botridii marginali sono dati 
da un 3olco circulare, abbas tanza profondo, che corre a guisa di 
un meridiano attorno alio scolice globoso, passando per 1 'apice, 
e diviciuendalo come in due emisfrri, uno :"estro a 1 'altro sinistro," 
thus evidently ignoring the fact that Luehe had already (1899 : 25) 
performed the service for students of the group, as pointed out with 
justifiable emphasis by the latter (Ariola, 1901 : 414). 

The bothria in this species ire sac-like structures, formed 
phylogenetically, as indicated in the generic diagnosis, by the 



— "1 

£15 



rolling together of their edges or "walls" and the fusion of the 
latter for moat of their extent "in 3uoh a manner that only a 
■nail lateral (dorsoventral) opening in the region of the apex 
leads into the interior of the spacious, hollow organ of attach- 
ment." The 3ize and shape of the opening itself varies consiera- 
bly in preverved material. It may be 30 small (Fig. 73) a3 to be 
seen only on very close examination or in sections, or comparatively 
large (Fig. 77), depending on the stage of contraction or enlarge- 
ment of the bothria when the individual is fixed or preserved. 
During life it may be seen to undergo 3uch variations in size while 
the whole scolex is being elongated and retracted during the char- 
acteristic sucking movements. Rudolphi (1813 : 477) correctly 
described the bothria as " ... oblonga profunda et magna in vivis; 
in mortuis bothrii ostium parvum anticum adesse viietur." In 
lateral view (Fig. 74) the bothria are seen to be more sharply oval 
or even conical in outline, as is consequently the whole scolex, 
owing to the fact that .the dorsoventral diameter of the lumen of 
each is much greater in the posterior half than in its anterior 
half. It will also be noted more clearly from this aspect that 
the hinder borders of the bothria project a considerable distance 
beyond the true anterior end of the strobila, so]that the length 
of the 3colex is not that of the bothrium, as many writers have 
evidently taken it to be, but as far as can be determined from 
external views, more nearly that of the marginal sulcus, plus an 
extension of the same to the tip of the 3colex, or, where the latter 
Is retracted, to the anterior border of the labia. The breadth of 
the scolex was taken for the sake of convenience to be that of the 



216 

bothrium, since there is very little difference between the two 
in this regard. 

The two apertures of the bothria are united over the tip of 
the acolex by a saddle- shaped groove, the edges of which are aome- 
what swollen so as to form lip-like structures. This groove has 
been described and figured for 0. eras sl eeps by Molin (1831 : 335, 
Fig. 2, Tab. V) and Ariola (1900 : 397, Fig. 17, Taf . VIII ) and 
figured by Linton (1310b : Fig. 257, PI. 34), but it does not 
appear either in the figures given by Wagener (1854 : Fig. 75, 
Taf. 7; 1857 : Fig. 6, PI. II) or that by Jonnstone (1309 : 37, 
Fig. 14). It is present in all of the writer's material even to 
the youngest, but in a few cases the tip of the groove, that is, 
the extreme tip of the scolex is so prominent as to more or less 
obliterate the lips (Fig. 77). It is al30 to be noted that the 
lateral grooves separating the bothria do not pass thru these lips, 
as nicely indicated in Ariola' 3 figure but erroneously described 
(p. 388) as "passando per l'apice", and as further figured but in 
the same relation by Johnstone. Wagener's Fig. 75 and Linton's 
Fig. 366 also give the erroneous impression that this groove passes 
right over the tip of the 3Colex. Molin (18S1 : 235), while 
giving a somewhat confused description (vide supra) of the rela- 
tions between the saddle- shaped structure, which he figures as in- 
cluding the apertures of the bothria more posteriorly, and the 
lateral grooves, saye that he saw in the apex an aperture which not 
only ended blindly but which Was bounded by two eminences, simula- 
ting lips. This may have been due to extreme contraction of the 
tip of the 3colex between the lips of this groove. It will be 



217 

recalled that F. 3. Leuckart (1819 : 46) says in this connection 
that "An dem Kopfende 1st eine kleine Vertiefung in der Blitte; die 
vcn den beiden sich hier verainigenden Randfurchen "aerruhrt, wo- 
durch ihre Rander etwas erhabener werden. Die Gruochen sind kaum 
von der Grc-33e eine3 Nadelknopf chens und tief in Kopfe, so das3 sa 
fast 3cheinen k'6nnte, als waren sie wahre 03cula," but his Fig. 33, 
very good in other respects, does not do justice to his description 
cf these terminal structures. Cf. also Loennberg's (1893 : 15-17) 
B. negl ectus, the figure for the scolex of which looks very much 
like 3. cra 33iceps . 

There is no neck in this species, segmentation beginning 
immediately behind the 3colex (Fig. 76) and being' complete thruout 
the strobila, which characters are also given by Luehe (1899 : 44) 
for the genus: "Aus3ere Gliederung vollkommen, ein gegliederter 
Hals fehlt." As regards this quotation, it would appear that the 
"gegliederter" is either superfluous or a lapsus calami for "unge- 
gliederter." The anterior border of the first segment , a greater 
part of which is obscured by the hinder edges of the bothria, is 
constantly somewhat narrower than the latter, but its posterior 
border i 3 usually about the same width even in such contracted 
specimens (Fig. 73). Its outline is somewhat trapezoidal, while 
its length i3 3lightly greater than that of the segment immediately 
following. The breadth of this first segment varies anteriorly 
from 0.40 to 0.93mm. and posteriorly from 0.35 to 1.16, — Linton's 
measurements are 0.78 and 1.07, respectively. Following this the 
segments are closely 3et, five to 3ix times as broad as long, while 
their somewhat thickened posterior borders protrude on either 3ide 



318 

(at well as dorsoventrally) 30 as to give the 3trobila a serrate 
appearance (Fig. 76). It is here that the formation of new pro- 
glottides takes place by the subdivision of preexisting segments. 
This serrate appearance is al30 present in the posterior part of 
the strobila where the proglottides are quadrate to t iriee as long 
as broad. The measurements of the first proglottis showing eggs 
in the expanded end of the uterus (uterus-sac), in a fairly re- 
laxed strobila (Fig. ), was 0.50mm. long by 0.92 broad, while 
one further back where the uterine cavity was 0.31 x 0.48mm. was 
1.31mm. long by 0.83 broad. These measurements are, however, of 
only relative value, since another strobila of the same age but 
contracted during fixation might show the same regions more like 
those farther ahead and thus, in alcoholic, specimens evidently 
younger. But posteriorly, however, each serration doe3 not nec- 
essarily define the posterior border of a proglottis. This is 
due to the presence of spurious articulations, possibly included 
in Wagner's "articulatio spuria" (vide infra). These are furrows 
which arise laterally, where they do not stand out as distinctly, 
however, s the true iiosterior borders of the proglottides, but 
do not pass to the median line. They are not present in all of the 
posterior proglottides nor are they symmetrically arranged. In 
the following excerpt from his more complete diagnosis Rudolphi 
(1819 : 477) did not refer to these structures: 

"Articuli breves, margine posteriore incra33ato 
utrinque exstante, quo corpus serratum fiat. Arti- 
culi ceterum inaequales, ut passim augustiores et 
longiores intercurrant . " 



219 

while F. S. Leuokart said only that 

"Die ersten Glieder am Kopfe 3chmaler als die 
Ubrigen, dann folgen fast gleichbreite , die 
letzte Halfte der Glieder breiter als lang, mit 
deutlichen, weissen Ovarien." 
which refers to "der beschreibene nicht ganze Wurm . . . i-i/2" 
lang." Diesing (1863 : 236) described the strobila aa 

" ... ellipticum, articuli3 ad medium usque increfcscent- 
ibus, inde desorescentibus, marginalibus posticis 
utrinque prominentibus , articulo 3ingulo plica 
transversali diviso ... " 
which latter refers obviously to T^agener's "articulo 3puria"; 
while it is also to be 3een that, a3 regards the shape of the 
strobila, he was dealing with much contracted 3pecimen3, the 
length being cited as ranging from one and a half times to two 
inches. Ariola (1900 : 397) said: 

"Strobila anteriormente assai piu 3tretto 
dello 3colice, a guisa di peduncolo; le primt 
proglottid! 3ono rettangulari , strette, ma 
rapidamente si allargano; raggiunta la ma33ima 
dimensione, la conservano sino all ultimo tratto 
del corpo, dove nuovamente si restringon*. Le 
proglottidi mature hanno angula posteriori arpena 
visibili; le ultime pre3entano forma trapezoidale . " 
and Johnstone (1909 : 89) stated that 

"The posterior j roglottiies are much broader (in 
the transverse axis of the 3trobila) than they are 



220 



long (in the longitudinal axis of the strobila); 
and. their anterior extremities are narrower than 
the posterior one3, so that the edge of the stro- 
bila appears to be serrated. Secondary segmenta- 
tion of the proglottis often occurs." 

In fine, Wagener, Diesing, and Johnstone are, to the 
writer's knowledge, the only workers who have referred to this 
spurious articulation or subdivision of ths segments into false 
secondary segments, — although Luehe (1902 : 833) repeated the 
statements of the first two authors. Furthermore Wagener did not 
figure the adult strobila of the species to show the structures in 
question, but in the legend for Fig. 79, Taf . 7 of Dibothrium 
heteropleurum, — now, Amph icot yle heteropleura (Diesing) — says 
only that "Man sieht die articulo spuria, welche die echten 
Glieder, wie bei Dibothrium orassioeps . in der Mitte theilt", and 
further, as regards the difference in structure of the sides of 
this species, "Der Schein entsteht durch die noch dichtere Zu- 
sammendrangung der Falten der wahren und falschen Glieder auf der 
concaven Seite." In his legend (p. 61) for Fig. 5, the egg of 
ff. orassioeps , he also said: "Jedes Glied hat in der Mitte eine 
Falte, die ihm das Ansehen giebt, als bestunde ss au3 zwei Gliedern. 
Thus there is reason to believe that for this species no one 
(apart from Linton's Fig. 233) has as yet described nor figured 
what the writer here calls spurious articulations, but that these 
workers were referring to the secondary division of the segments 
of the anterior end of the strobila which proceeds in the manner 



331 

described for B. soorpii et al, altho not 30 clearly (Figs, 76 and 
73). This is borne out by the fact that the 3purious arti eulations 
described here never reach the median line of the strobila, much 
less pass completely acroas it a3 do the true posterior borders of 
the proglottides (Fig. 73). In one moderately relaxed 3trobila the 
first segment showing 3purious articulations appeared 11.7mm. from 
the tip of the scolex, while in another which was quite contracted, 
especially anteriorly, 4.8mm. In the former case the next two 
pairs of these structures, — and all of these in question happened 
to be bilaterially symmetrically situated, — appeared in the fourth 
and thirteenth segments following. 

The following table gives various external measurements of 
specimens in alcohol, which may be of use for comparison: 



Number of specimen 


304.1 


304.2 


304 . 3 


204.4 


331.1 


259.1 








Little 






Length 


87mm . 


92mm . 


43mm . 


more than 


29mm 


72mm 








3colex 






Length of scolex 














(lateral view) 


0.87 


0.59 


0.46 


0.43 


0.63 


0.83 


Length of bothrium 


1.08 


0.77 


0.54 


0.64 


1.00 


1.01 


Breadth of scolex 














(bothrium) 


0.75 


0.57 


0.52 


0.53 


0.67 


0.90 


T:.ickne3s of same 


0.87 


0.64 


0.68 


0.58 


0.74 


1.21 


Breadth of Seg. I 


Much con- 


0.40 


0.37 




0.55 


0.92 


(anteriorly) 


tracted 












Ditto, posteriorly 


Ditto 


0.53 


0.60 


0.53 


0.74 


1.16 


Thickness, posteriorly 


0.37 


0.38 


0.24 


0.37 


0.52 




Greatest breadth 














Anterior part of prog 


1.01 


0.83 


1.06 




1.30 





Posterior " " ■ 1.11 1.04 1.16 1.48 1.38 



333 

None of ths above six specimens were considered to be complete 
posteriorly. The posterior proglottis will be dealt with below 
under the excretory system. 

Posteriorly the uteru3-aacs appear as a series of gradually 
enlarging, dark punctations, a3 desdribed below, not 30 pronounced, 
however, as in B. scor pii . 

The cuticula varies in thickness from 2 to 5^ , the most 
common measurement being about 2.Syu . Resting on a distinct base- 
ment membrane, -veil shown after the use of Mallory'3 stain, it is 
divided into two strata of equal thickness by a granular layer, 
the components of which seem to be relate! to the bases of the 
stout, somewhat club-shaped bristles or "hairs" which constitute 
the outer moiety. While the inner stratum was found to be homo- 
geneous with the stains u.3ed, the outer showed two intensities of 
color, an inner lighter and an cuter darker. The former repre- 
sents the narrowed central end3 of the spindle- or club-3haped 
bristles, while the latter is determined by the veil-stained 
bodies of the bristles themselves. Linton (1301 : 473) said that 
"the cuticula is covered with minute spines", but Johnstone (1909 
: 39) said concerning these structures: "I can see nothing of this 
kind in the 3pecie3 before me." All over the scolex and in the 
form of a band on the posterior borders of the proglottides (Fig. 
82) thi 3e bristles become modified into stouter spinelets from two 
to three times as long and everywhere directed posteriorly, quite 
like those described elsewhere (Cooper, 1314b : 85) for Kap lobo- 
thriurn glob uli forme , but :mch longer relatively; thus indicating 
their function a3 accessory organs of attachment. The largest 



333 

3pinelets are in the middle of this band, those at the edges, that 
is in the antero-posterior direction, gradually merging in length 
into the bristles of the cuticula of the neighborhood. Furthermore 
they are arranged in the same manner- on the posterior borders of 
the spurious articulations and all the secondary segments situated 
in the anterior portion of the strobila. They were originally re- 
ferred to by Wagener (1354 : 5) and later by Die3ing (1863 : 236) 
("articulo aingulo ... postice ciliis instructo") , by Cohn (1902 : 
55) and by Luehe (1902 : 238, 247) who considered "dass es sich 
nicht um in die Cuticula eingesenkte 8tacheln hands It, wie bei dem 
Stachelkleide so vieler Distomen, sonde rn nur am Fortsatze der Cuti- 
cula, durchaus analog denjenigen, welche Looss an der bereits oben 
citierten Stelle fur Haemat oloe chus a_3p_er abgebildet hat.." 

The cubcuticula, about 30 ^ in thickness, consists of fairly 
elongated cells, the nuclei of which are situated at their central 
ends close to the vitelline follicles ; while their boundaries are 
difficult to ascertain, the whole layer thus being more of the 
nature of a syncitium. For about one third of their length immed- 
iately beneath the cuticular the cytoplasm becomes broken up into 
a number of more or les3 parallel processes which stand out in dis- 
tinct contrast with the deeper inrer ends of thecells especially in 
transverse sections. 

The parenchyma, everywhere encroached upon by the voluminous 
reproductive organs, is in the form of a comparatively open reticulum 
showing no features of special inter. st. It is naturally most abun- 
dant in the posterior flared end3 of the proglottides. In small 
3trobila it is more compact in structure and has relatively more 



234 

nuclei in its me3hes. Distinct spaces, formerly occupied by- 
calcareous bodies, such as are readily and distinctly 3een in the 
parenchyma of B. scorpi i , were found neither in the scolex nor in 
the strobila; nor were these structures noticed in liv.'.ng material. 

The musculature is composed of the typical three set3 of 
fibres, interf erred with in the usual manner by the large reproduc- 
tive organs and their external openings. The sagittal and coronal 
3erie3 are only moderately developed, while the longitudinal series 
if about lOjjL in thickness and situated within the coronal series. 
Its fibres are arranged in bundles of irregular shape (in cross- 
section) and width but of this uniform thickness, excepting where 
they are naturally much flattened out dorsally and ventrally by the 
distended uterus-sac. Otherwise they are continuous from joint to 
joint. A very weakly developed series of outer longitudinal muscles 
is also present while the muscles of the posterior border of the 
proglottis (vide Luehe 1397a) are poorly developed, in fact even 
less so than in Bothriocephalus, s.str. The cuticular musculature 
is typical. 

In the ecolex the coronal fibres are better developed than 
the sagittal ones and pas3 around the bothrium closer to its lun.en 
than to its external surface, while the latter are mostly confined 
to the region between the bothria. The inner longitudinal muscles 
of the strobila pa33 forward into the scolex, dividing as they meet 
the lumen of the bothria to pa33 around them and attach themselves 
to the margins of the apertures. They are thus directed somewhat 
obliquely a3 shown in Johnstone's Fig. 18 and|described as " ... 
running irregularly, probably obliquely, round the walls of the 



325 



bpthrlum. These no doubt function as constrictors of the latter." 
A few pass on forward to the tip of the a 00 lex to assist in activa- 
ting that region. Between the bothira, however, they were found to 
be separated into dorsal and ventral layers as in the 3trobila, and 
not united into a single coronal band as shown by Johnstone. The 
bothrial sphincter (Fig. 76) is a powerful bundle of fibres, about 
0.07mm. in transverse section surrounding the aperture close to its 
cuticula. In transverse sections of the 3Colex it appears as a 
deeply staining ma33 on each side of the opening, also shown in 
Johnstone's Fig. 15. As it crosses the aperture anteriorly it 
becomes greatly attenuate!, nrhich fact with its comparatively great 
size at ths 3ides and posteriorly accounts for the almost complete 
disappearance of the aperture in many adult, preserved scolices 
owing to the po?;erful contr ction of this muscle from behind forward 
thus diminishing the opening towards the tip of the scolex. From 
their arrangement it i3 to be seen that this sphincter, evidently a 
modified group of coronal fibres, and the longitudinal muscles in 
the scolex play a more important role in the movements of the 
bothria than do the other groups. On account of the oblique course 
of the longitudinal fibres they would evidently act in diminishing 
the size of the lumen of the bothrium as ..ell as vould the circular 
(coronal) fibres of the latter. 

The nerv^ous system consist 3 of two longitudinal 3trands 
which enlarge in to tip of the scolex to form two some~;hat elongated 
ganglia, united by only a few fibres but sending out conrparatively 
large nerves to the bothria. In the s^robila the chief strands, 
each from 15 to 30 in diameter, are situated ventrally in the 



326 

medullary parenchyma just within the longitudinal muscles and 
from one-fifth to one-quarter the width of the strobila from its 
lateral margins (Fig. 80). About half way along the 3colex the 
3trands are about 80 ^ in diameter, while the somewhat smaller 
ganglia are close together about 0.15mm. from the summit. In other 
words the chief strands enlarge and diverge gradually until the 
equatorial region of the scolex is reached and then diminish in 
size as they converge to form the ganglia. A pair of prominent 
nerves is sent forward on each 3ide to supply the saddle- shaped 
groove described above. In young strobilae the nerve strands are 
situated midway between the dorsal and ventral surfaces, and not 
ventrally . 

The excretory system consists of a pair of longitudinal 
vessels, situated ventrally, that is in the same frontal plane as 
the chief nerve strands, each vessel being in the anterior end of 
the strobila about half way between the nerve strand and the median 
rows of reproductive rudiments. These vessels break up in a very 
irregular manner into extremely elongated loops, so that for con- 
siderable stretches four vessels will appear while again the 
branchings will be so numerous as to make it very difficult to de- 
cide, on looking at a transverse section, which are the main chan- 
nels (Fig. 76). In other individuals four vessels appear, so that 
we must conclude that the pair ju3t mentioned represent the latter 
fused at times but separated again to form the loops. But whether 
these four vessles represent the typical four of other orders it 
was found impossible to decide, excepting from comparisons with 
other sp-. cies of this order. These main vessels may continue back 



227 

into the ripe joints cl03e alongside the uterus- sacs, but they 
usually break up into a very diffuse reticulum throughout the 
medullary parenchyma in the region where the openings of the cirrus 
and vagina pierce the cuticula in development. Behind this region 
it was found impossible to trace the main vessel 3 with satisfaction. 
The system usually passes into the seolex as two vessels, but soon 
breaks up into an elaborate reticulum which ramifies between the 
bothria and throughout their walls. These branches are shown in 
Johnstone's Fig. 15. A3 regards the conditions of the excretory 
system in the extreme posterior end of the strobila, the material 
at hand permits of only negative conclusions. In the youngest 
3trobilae, such as that shown in Fig. 77, the vessels converge 
posteriorly to open into a notch in the cuticula, there being no 
definite pulsatile vesicle such a3 is present in plerocercoids of 
the genus p roteoce::halu3 , for instance. From this and the further 
fact that Wagener (1357 : 93) showed (Fig. 0, PI. II) the main 
vessels in a very small strobila, which he examined while it was 
alive, passing separately to the outside, we are led to conclude 
that the vesicle, if every present, must have been situated in the 
walls of an enveloping cyst and disappeared with the latter as in 
the Trypanorhy^cha or the Cyclophyliidea . This seems to have oeen 
Wagener' s idea of the situation when under his Fig. 65 (1854 : 68) 
he said: "Man sieht keinen pul3irenden Schlauch am spitzen Schwanz- 
ende . Es mu3s dies Thier auf ahnliche Weise entstanden sein, wie 
das in Fig. 74 dargestellte , " and Fig. 74 i 3 that of "Dibothrium 
(Belcnes?)" from Soyllium can! cula enclosed in a cyst in the walls 
of which "man sieht der Gefasse der 0e3todenblase . " 



338 

The earliest refer nee to the genitalia of C. ora 3 sleeps 
was by Rudolphi (1319 : 477) where he said: 

"Ova vel ovalia vel ovata, forsan secundun 
majorem maturitatis gradum. A 3. punctato diver- 
sissinms, licet ovaria lateralia fusce^Lscant , 3ed 
haec ipsa etiam in B. orasslcipite quam in B. pune- 
tato majora sunt." 

The structures called ovaria were evidently the uteru3-sacs. 
F. S. Leuckart (1819 : 46) described the reproductive organs of 
his 3. pilul a as follows: 

" ... die letzte Ealfte der C-lieier breiter als 
lang, mit deutlichen, weissen ovarien. An den 
uteren Gliedern sieht man oberhalS jedes Eier- 
stockes einen 7/asserhellen Funct, wahrscheinlich 
Oeffnung fur das mannliche Zeugungsglied . " 
From a comparison of this with his description and figure of pos- 
terior proglottides of B. scorp ii , it is evident that he too was 
dealing with the uteri and their openings respectively. He also 
referred to " ... den schwarzen Puncten des Korpers, die Rudolphi 
fur Ovarien gehalten" of Recli ' s worm, which Rudolphi called 
(1310 : 67) Bo t hri o cephalus Gadi merlucp_ii_ and placed in his 
"Species dubiae." Wagener (1854a : 31) said that 

"Die Eier Haufen 3ich in obersten Theile der 
Glieder an. Der Dotterstock verzweigt sich uber 
das ganze Glied und liegt overhalfc des vescicules 
tr:-.nsparentes van Beneden. Die Geschiechtsof fnung 
ist in der Mitte und lateral." 



229 

Dies^.ing (1883 : 236) placed the "Aperturae genitalium laterales 
in linea mediana." Ariola (1896 : 255-366) gave the first com- 
prehensive description of the reproductive organs in the following 
words : 

"Tuttaria aul corpo si osservano macchie acure 
molto sporgenti , con3tituite della massa di uova. 
Tali rilievi non sono propriamente , nella linea 
mediana, ma collocati a destra o a sinistra di 
essa, formando in tal modo une striecia a zig-zag. 

L 'aperture geni tale msschile sbocca sulla 
faccia dorsale, e sulle oppoata si apre l'utero. 

In alcune proglottid! l'ovario e bilobo, la 
uova sono ellisaoldall e mancano di operculo." 
Luehe (18SS : 42-44) in defining the characters of the genus gave 
the general features of the genitalia, while Ariola (1900 : 397) 
enlarged hi 3 own 18S6 description: "Ovario con numerose uova, 
talora bilobo; uova ellissoidali aventi nel iiametro longitudinale 
|^ 57 e nel traaversale 32 ... " Braun (1900) reviewed the litera- 
ture on the genua and apecies up to date, and Volz (1900) diacussed 
the reproductive organs of the apeciea as compared to those of hi a 
B. apiralicepe and the position of the openings in connection with 
brief remarks on the j hylogeny of the genu3 Bo t h r i o c ep ha lu s s.lat. 
As regards his own specimens Linton (1901 : 473) said that "Pos- 
terior segments show rudiments only of the reproductive organs, but 
no indication of external genital openings." And later Johnstone 
(l.o. : 89) remarked that "the genital openings are in the middle 
line of the proglottides but near the anterior borders of the 



330 

latter," referring evidently, as will be seen later, to the uterine 
openings only. 

The rudiments of the reproductive organs appear about 
three millimetres from the tip of the scolex as aggregations of 
nucleu that can just be discerned in to to mounts (Fig. 73). About 
three millimetres farther posteriorly in moderately contracted 
(such as would be obtained if no special care were taken during 
the fixation of the material) older strobilas the cirrus and 
vagina are seen to be just piercing the dorsal surface. Before 
this region is reached, however, the common rudiment, at first 
circular and then elongated oval in outline, differentiates into 
a more anterior portion, the rudiment of the whole uterus, a more 
posterior less elongate.! part, the beginnings of the cirrus-pouch 
and vagina, and a third, connecting the other two near the hinder 
edge of the proglottis, the nuclear aggregation that will develop 
into the ovaries and the organs of the interovarial space (Fig. 79). 
As mentioned above in the specific diagnosis, the first two of 
these rudiments alternate irregularly from 3ide to side as do the 
corresponding adult structures. At the same time the testes and 
vitelline glands are developing in the medullary and cortical por- 
tions of the parenchyma, respectively. 

A distinct genital sinus or cloaca, the opening of which 
is usually almost circular in outline, is present (Fig. 83). It 
varies from 9.05 to 0.09mm. in diameter and is situated, as a*;ove 
noted, nearly in the median line, dor sally, and from three-fourths 
to one-half the length of the proglottis from its anterior border, 
usually just posterior to the 3vurious articulations when they are 



— I 

331 

present. At the bottom of this sinus there is a secondary cloaca 
( "Geschlechtstasche " or "Ductus hermaphroditicus") , also circular 
in outline, from 15 to 35^ in diameter, and into it open the cirrus 
and vagina quite close together, the latter immediately behind the 
former. This secondary 3inus is best 3een in sagittal sections 
(Fig. 82). The genital pore (the opening of the main sinus) is 
elevated slightly above the general dorsal surface of the immediate 
neighborhood, thus appearing as a low cone or crater. No sphincters 
were found to control the openings of either of these sinuses but 
the cuticula of the floor of the larger or outer was seen to be 
modified into coarse, low, rounded and close 3et papillae which 
are evidently of special inportance during copulation. As regards 
the latter it was concluded that these papillae would serve to 
temporarily fasten the structure into the primary sinus of another 
proglottis, when it is possibly everted with the sirrus. Nothing 
was observed on copulation in this species during life nor were 
any case 3 of protruded cirrus met with in the material at hand. 

All of the proximal portions of the reproductive organs, 
excepting the vitelline follicles, are located in the medullary 
parenchyma, although the much distended uteru3-sac, originally in 
the latter, extends almost to the cuticula on both the dorsal and 
ventral surfaces. Fig. 81 shows their- arrangement in toto. 

T he testes are closely arranged in the medullary parenchyma 
in two lateral fields, each bounded laterally by the junctions of 
the dorsal and ventral layers of longitudinal muscles and medially 
by the other reproductive organs (excepting the vitelline glands) 
which occupy in the quadrate proglottides about the middle one-third 



332 

of the transverse diameter of the strobila and are contiguous from 
joint to joint. In the quite mature elongated proglottides the 
testes are ellipsoidal in shape, averting 0.135mm. in length by 
0.040 in diameter, the cro33- section being usually about circular 
in outline. In younger joints and in all those of much contracted 
strobilae the testes are nearly spherical in shape, measuring about 
60 p. in diameter, or often slightly longer than broad. They are 
arranged in a single layer in the medulla, the -.'/hole dorsoventral 
diameter of which they occupy, and are continuous from proglottis 
to proglottis. From 2 to 4 appear in each lateral field in trans- 
verse sections, from 5 to 7 are 3een in sagittal sections between 
the posterior borders of consecutive proglottides, while, so far 
as could be determined from coronal series directly, the number i3 
from 20 to 25. Thus each proglottis contains from 40 to 50 testes. 

The vas deferens forms a wedge-shaped ma;s of closely ar- 
ranged coils, extending forward immediately ahead of the cirrus- 
pouch and alongside the uterus- sac for about two- thirds of its 
length (Fig. 81). In proglottides in which the latter is yet 
comparatively small the vas deferens may pass forward as far as 
its anterior end. In either case it forms with the cirrus-pouch 
a mass which alternates from right to left with the uterus- sac. 
When distended with sperms the duct averages about 30 in diameter; 
but just before it enters the cirrus- sac anterodorsally it narrows 
down to 5 f*. . Immediately within the wall of the latter it often 
enlarges again to form a thin- walled functional vesicula 3eminalis, 
or perhaps more correctly ductus e jaculatorius , from 13 to 23(jl in 
diameter. After one or two short turns it diminishes again to about 



233 

8 j^- and then passes on as the cirrus proper. While the ;.roximal 
portions of the iuot do not pa33 in any definite direction, the 
latter is situated for most of its length in the longitudinal axis 
of the pouch and is about 0.10mm. long. About 20 to 35 p. at its 
middle, it is lined with a cuticula, 10 p. thick, which is cleft 
but net armed with bristles of any kind. 

The cirrus.sac (Fig. 83) situated immediately behind the 
uterus-sac or lateral to its rosterior end, is elliptical to slight- 
ly oval in outline, and measures .128-0 .132mm . long, .087-0 .116mm. 
Tide and 0.098-0.116 deep. The longitudinal axis is directed 
anterodorsally from the genital 3inus ana to the right or left, 
according as it alternates with the uterus-sac. The proximal 
one- third of the contents of the pouch consists of loose parenchyma- 
tous tissue with a few muscle fibres surrounding the ducted ejacu- 
latorius, while the distal two-thirds, that part which accommodates 
the cirrus proper, is supplied mostly with muscles which actuate 
the latter. Large fibres proceed somewhat obliquely from the wall 
towards the proximal pole of the sac to become broken up or frayed 
before they are attached to the cirrus tangentially, so as to give 
the appearance in frontal sections of the latter being surrounded 
by a comparatively heavy layer of fine lightly staining circular 
fibres. A few of the fibres closest to the cuticula of the cirrus 
were considered to be true circular fibres; but no longitudinal 
fibres were seen. The wall of the cirrus- sac is from 2 to 3 ^ 
thick and is made up of very fine closely matted fibres, the direc- 
tion of which could not be determined with satisfaction. The sac 
lies freely in the parenchyma of the region and is not connected 



234 

"by any special muscles to the dorsal or ventral body- walls; nor 
are the body muscles attached to it as in some c3.3todes. The 
layers of the latter are simply pierced and the fibres turned aside 
in evidently a passive manner. 

The opening of the vagina is close behind that of the cirrus 
at the bottom of the secondary genital sinus, or as it has been 
called by Fuhrrnann, "ductus hermaphrocti ticus " (Fig. 82). From this 
point the duct courses ventro-posteriorly in the mid-line and then 
parallel to the dorsal surfaoe of the proglottis until it reaches 
the ovarian isthmus, above which it makes a few turns and quickly 
diminishes from 20^ in diameter half 7/ay along its course to 10^ . 
It then dips farther down into the genital , 3pace , often enlarging 
slightly as it does, and soon joins the oviduct at an enlargement 
of the latter situated a 3hort distance behind the oocapt. 
Throughout its length it is lined with a ragged or pseudociliated 
cuticula and surrounded by radially arranged nuclei connected with 
the cuticula by cytoplasmic strands like those described by the 
writer for H. giob uli forme (i.e. : 105) and considered to be 
possibly extruded nuclei of the original epithelium as well as the 
myoblastic nuclei of circular fibres, a ljjyer of which surrounds the 
duct. There is no vaginal sphincter. 

In his generic diagnosis Luehe said that the receptaculum 
3eminis is small and in his description of the family, Ptychobo- 
thriidae (1902 : 327) that when present it is "in Gestalt eines 
kleinen Blindsackchens ausgebildet, welches parallel neben dem 
Fndabschnitte des Oviduktes liegt and mit der Vagina unmittelbar 
vor deren Vereinigung mit dem Ovidukt in Verbindung stent." In the 



235 

sections at hand, however, it is a comparatively large structure 
and very difficult to orient in sections made in any direction. 
It is in the form of a thin-walled sack about 80x20 j/l , '.vrapped 
somewhat spirally around the dorsal wall of the above-mentioned en- 
largement of the oviduct and opening by an aperture equal to its 
whole diameter into the vagina just at its juncture with this 
vestibule. But since the vagina constantly constricts a second 
time to a diameter of about 8y^ before entering the latter, one 
gets the impression of the receptaculum seminis being a diverti- 
culum cf the oviduct rather than of the vagina. Fig. 83, of four 
consecutive sections of a transverse series, showing the union of 
these ducts, will give a better idea, perhaps, of the nature\of 
the seminal receptacle. 

In mature proglottides the ovary (Fig. SO) is a bilobed 
structure situated in the median line, close to the posterior 
border of the proglottis and immediately ahead of the uterus-sac of 
the proglottis following, where the latter is much distended with 
eggs (Fig. 81). In toto mounts the lobes seem to be quite separate 
from each other and apparently unconnected, but in sections the 
isthmus is easily made out. It occupies the ventral half of the 
medulla while the wings or lobes extend completely across the space 
betweenthe layers of longitudinal body muscles. The lobes are 
about 0.37mm. long by 0.13 wide, while the isthmus is .06-0 .08mm . 
in anteroposterior diameter. These proportions are, however, much 
different in such contracted strobilae or in proglottides in which 
the uterus-sac is distended with eggs. In both instances the ovary 
is very much flattened anteroposterior^ and, in the latter case, 



235 

all but obliterated, as shown in Luehe's Fig* 8 (1902 : 32S). The 
ova from the isthmus where they are ready to be passed on by the 
oooapt, are elliptical to oval in outline in 3ection3 and measure 
on the average 13 x 10 p., their nuclei being about 9f*. in diameter. 

The oooapt j situated in the median line at the posterior 
border of the ovarian isthmus, somewhat dor sally, is a spherical 
to ovoid muscular organ, about 30|A- in diameter (Fig. ). Imme- 
iiately behind it the oviduct constricts to a diameter of only 
to 10 jl and then passes on posteriorly and ventrally either to the 
right or to the left, gradually enlarging until the above-mentioned 
vestibule is reached, when the diameter is 25 to Z0\h . The latter 
enlargement does not seem to be a direct continuation of the ovi- 
duct but a more or less separate thin-walled structure, — the 
■vails of the oviduct up to this point being comparatively thick 
(Fig. 33) — into which the oviduct opens by a slightly elongated 
aperture. While the wall of the first portion of the oviduct is 
quite thick, comparatively speaking, and consists of more or less 
cubical ciliated cells with somewhat indefinite boundaries, — or- 
dinarily they stain very densely, — the vestibule has, like the 
receptaculum seminis, a thin wall from which only a few scattered 
nuclei protrude into the lumen. The oviduct continues posterio- 
laterally and ventrally from one cor&er of the vestibule — that 
with which the vagina is usually connected, — as a tube quickly 
diminishing from 15 to 10 p. in diameter and lined with a ciliated 
epithelium with prominent nuclei but no distinct cell-boundaries. 
Close to the anterior wall of the uterus-sac of the next proglottis 
it turns upward sharply and at about the middle of the dorsoventral 



2 37 

Hameter of the medulla takes on the vitelline duct. It then skirts 
the uterus-sac, just mentioned, as it passes to the opposite side 
of the generative 3pace and slightly forward to soon become sur- 
rounded by the shell-gland. 

The vitelline duct at Ita union with the oviduct has a 
diameter of 8fA, but just beyond this, in the direction of the 
follicles, it 30on enlarges to form a somewhat irregular vitelline 
reservoir which when filled with yolk may attain a diameter of 
30|A# . Its general course is towards the opposite side of the 
generative apace almost parallel to either surface of the body; 
but beyond this it could not be traced with satisfaction. 

The vitelline follicles fill up almost the -"hole of the 
cortical parenchyma from the layer of longitudinal body muscles to 
the nuclei of the subcuticula, the thickness of the stratum aver- 
aging O.oSmm. (Fig. 80). They form a continuous layer around 
the margins of the proglottides (in transverse sections) and also 
from proglottis to proglottis, as mentioned above, even extending 
well into the posterior borders. They are not arranged in lateral 
fields, but are interrupted only where the uterus-sac and genital 
sinus pierce the body-wall, or in the former case greatly press 
against the latter. The individual follicles attain a size of 50/u. 
long, 30pi nride and 50(a- deep (the thickness of the -"hole layer), 
and are very closely crowded together. The number in cross-sections 
of the proglottis averages 55 and in sagittal sections 13, thus 
making the average total number for each proglottis 715. 

The shell-gland is situated in the dorsal portion of the 
genital space, that part of the oviduct showing the connections 



338 

being almost horizontal in position and about 18^ in diameter, that 
is, a little larger than the oviduct behind that region. The indi- 
vidual cells of the gland are much attenuated, closely arranged 
and have their nuclei situated in their slightly enlarged distal 
ends. Their connections with the oviduct give the wall of the 
latter a honeycombed appearance when it is seen in longitudinal 
section . 

Beyond this region the oviduct gradually enlarges as it 
passes above the ovarian i3thmus to become the uterine tube, the 
coils of which are accommodated opposite the cirrus pouch just be- 
hind the uterus-3ac. As it proceeds Its wall get3 thinner, the 
nuclei protrude more and more into the lumen until many of them are 
evidently lost. It is noteworthy that the uterine tube in many 
cases as well a3 the uterus-sac especially in younger proglottides, 
alternates irregularly from right to left according as the cirrus 
and vaa deferens do. The3e three structures are, intact, fitted 
very nicely into the space between the uterus- sac ahead and the 
ovarian isthmus behind. 

The uterus- sac is elliptical in outline, has its longitu- 
dinal axis directed anteropo3teriorly , and i3 situated in the an- 
terior half of the proglottis where in very mature segments it 
occupies almost the whole of the medullary region, or to be more 
precise, the middle three- fifths of the Hamrter of the proglottis, 
its anterior and extending forward close to the ovarium of the pro- 
glottis immediately ahead (Fig. 81). Luehe (1902a : 32S) figured 
the uterus as, to use his own words, " ... in der Regel eine ge- 
r'aumige Uterushbhie bildend , welche die ubrigen Geni talorgane , ohne 



239 

dass freilich deren Plickbildung eintritt, buchstablich an die Wand 
drangen kann, indera die ganze Proglottis in reifen Proglottiden 
vielfach als ein einziger sackformiger Eibehalter mit verbal tnisS- 
nas^g 3ehr diinnen Wandungen erscheint." But Buch a degree of re- 
striction of the other genitalis was seen only in a few of the 
ripe proglottides of strobilas much contracted longitudinally. 
Here the largest uterus- sac measured 0.3mm. wide by 0.57 long, 
while the width of the proglottis in question was, at the posterior 
borders of the spurious articulations, 1.57mm. In fairly relaxed 
3trobilas it increased in dimensions from 0.18 x 0.14mm., where the 
first eggs appeared in the lumen, to 0.87mm. long by 0.48 wide, 
where the proglottis was 0.80mm. wide at its middle, in the latter 
case, of course, pressing against the dorsal and ventral walls 
even as far as the cuticula. From a comparison of these measure- 
ments and the further fact that in the case of the former much con- 
tracted strobilae there often appeared behind the region showing 
the nearly obliterated genitalia a more relaxed one in which the 
relations of the uteru3-sac to the other organs was quit: as in the 
completely relaxed strobilas, one would be inclined to conclude 
that the characters of the family above quoted, .vould apply to this 
species only in the case of proglottides much contracted longitu- 
dinally. In the quadrate proglottides the smaller, that i3, 
younger sacs alterate irregularly from right to left, as do the 
uterine openings, and according as the cirrus pouch and the vas 
deferens in particular (on account of its above-mentioned position) 
occupy the opposite si:le3 of the proglottis. Externally, in alco- 
holic specimens, the uteri appear as a gradually enlarging series 



340 



of brown punotations caused by the contained eggs showing through 
the thinned body wall, aa pointed out originally by Rudolph! and 
other writers. 

The wall of the uterus consists of a thin membrane on 
the inside of which a very few scattered and somewhat flattened 
nuclei indicatelita original epithelial nature. In young proglot- 
tides, where no eggs are to be 3een in the small uterine cavities, 
the wall was found to be composed of an epithelium about 3 \l thick, 
showing prominent nuclei but no distinct cell-boundarie3 . Further- 
more in such early stages the lumina of the uterine ducts, develop- 
ing in the manner described by Young and Shaeffer, are not complete- 
ly formed nor in connection with the cavities of tho> 3acs, but 
the uterine apertures are prominent. In the first two or three 
sections of a 10 jjl coronal series, taken from the ventral surface, 
they appear as distinct somewhat elliptical apertures about 35^ in 
transverse diameter, but in the third or fourth section are closed, 
only to reopen as the cavity of the uterus- sac, thus showing that 
the membrane closing the aperture is only about 10 ^ thick. And 
this closed condition is maintained until the uterus-3ac attains 
the above-mentioned maximum 3ize and becomes greatly distended with 
eggs. Then the functional opening is established by the rupture 
of the membrane which has meanwhile reached a length of 0.043 — 
0.058mm. by a width of 0.034-0.045, it3 elliptical outline having 
been retained. The opening does not become as regular in outline, 
however, as the membrane, for the latter remains around the rim as 
ragged processes, which render the determination of the exact loca- 
tion of the aperture in toto mounts a matter of no little difficulty. 



241 



The uterus opening is surrounded by a series of radiating cells 
like those of the opening of B. soorpii described above. 

The fresh eggs examined in saline solution were found to 
be elliptical to ovoid in shape, 75 by 4:0^ in dimensions and pro- 
vided vith a thin very light brown shell having no operculum. The 
color waa so faint that it could be seen to advantage only when 
the eggs were in masses or in the uterus- sac. Ariola (1900 : 397) 
gave the measurements of the eggs of the European species as 87 x 
32|A- . The largest examined were immature, the contents consisting 
of large spherical cells like those shorn, by Wagener (1854a) in 
his Fig. 6, Taf . I, among which no traces of the hooks nor division 
into oncosphere nor mantle could be 3een. '"hen the scolices of 
the worm are still attached to the\wall of the intestine of the host 
between the mucous folds, they were round to often discharge many 
of their eggs from most of the posterior proglottides when the 
3fiolices were irritated with a blunt needle inorder to make them 
loosen their hold, which indicentally is a comparatively firm cne 
as one might gather from the structure of the seolex. 

A3 regards the life history of the species nothing definite 
was determined. 44 specimens of Merluccius bilinearis v«rere exam- 
ined at Woods Hole and at Harp swell, but no definite idea of a 
possible intermediate host was obtained. It was noticed, however, 
that when the intestine of the fish contained much grey chyle, pre- 
sumably the result of the digestion of small herring, — definitely 
ascertained at South Harp swell to be such in a few cases, and of 
the Blueback p Pomolobus aest ivalis (Hit chill ) , — no tape-worms of 
this species were present; but where amphipoda were found in the 



242 



stomach or the remains of such in the intestine the worm was plen- 
tiful. Furthermore, where nothing was found in either 3tomach or 
intestines, other than yellowish chyle in the latter, — as in most 
fish examined — indicating amphipod£ and other small crustaceans 
as food rather than small herring, the worm wa3 also common. All 
stages from the youngest strobilae, such as that shown in Fig. 77, 
to the oldest were found, but none nor any plerocercoids, if such 
is the nature of the larva, were met with in the course of the 
thorough iissection of the available stomach contents of the host, 
both fish and crustaceans. In a number of cases, nevertheless, 
only very young strobilas were found in the intestine of the host, 
thus pointing to possible sudden infections at different times. 
TTagener, who figured the youngest strobila, nothing much more than 
the scolex, that has yet been recorded, say3 nothing more concerning 
the life history than that, on account of the excretory vessels 
opening separately to the exterior in this very young specimen, 
there might possibly have been a veaicular appendage to the larva 
in the nature of an enveloping cyst comparable to that described and 
figured for "Dibothrium (Belones?) "from Scyllium canicula, con- 
cerning which he said (p. 45): "Vergleicht man diese Form von 
Cysticercus mit den vorigan, 30 ergiebt sieh, das der Unterschied 
nur in dem Auf hangebeutel sich findet, der Kopf und Blase verbindet" 
( Qyat i cercus fasciolaris Rud . ) 

A detailed description of the species is here given, not 
only because it is evidently the only one belonging to the genus, 
but because 3uch seems to be quite lacking from the European litera- 
ture, which male the determination of the species here a matter 



243 

attended with considerable uncertainty. The writer, however, 
considers that so far as the published accounts and reports of the 
species go, we must lock upon the form on this side of the Atlantic 
to be the same as the C . crassiceps of Europe . 

The material studied consisted of eight lots in the writer's 
collection from Merlucoius bi linearis a$4 above listed. 



244 

Subfamily 2. AMPHICOTYLINAE Luehe, 1902. 

Scolex with two typical, mostly not very deep bothria, which 
can nevertheless develop posterior, sucker-like portions. In an 
isolated case a pseudoscolex is substituted for the scolex. Ex- 
ternal segmentation insignificant, at times disappearing thru acces- 
ory wrinkling or folding of the surfaces of the proglottides. 
Opening of cirrus and vaginal marginal, irregularly alternating, 
with more or less strongly pronounced tendency to unilaterality. 
Uterus-opening median; uterus-sac always well developed. Coiling 
of vas deferens strongly expressed. 

Occurrence: In fishes. 

Type genus: Amphicotyle (Diesing, 1864) Ariola 1900, e.p. 
Luehe 1902. 



Genus 1. Abothrium van Beneden, char, emend. Luehe, 1899 . 



Taenia (part.) 


Auctorum. 




Rhytis (part.) 


Zeder, 


1803. 


Eothriocephalus (part.) 


Rudolphi, 


1809. 


Bothriocephalus (part.) 


Rudolphi, 


1819. 


Bothriocephalus (part.) 


Leuckart , 


1819. 


Bothriocephalus (part.) 


Dujardin, 


1845. 


Dibothrium (part.) 


Diesing, 


1850. 


Bothriocephalus (part.) 


Baird, 


1853. 


Dibothrium (part.) 


Diesing, 


1863. 



245 



Bo thriocephalus (part.) 


01s son , 


1867. 


Abothrium 


Beneden, 


1871. 


Abo thrium 


Lloniez , 


1881. 


Dibothrium (part*) 


Linton , 


}890. 


Abo thrium 


Loennberg , 


1891. 


Bothriocephalus (part.) 


Matz , 


1892. 


Ro thriotaenia inapt. ^ 


Ariola, 


1896. 


Eothrio taenia (part.) 


Riggenbach,1896. 


Abothrium 


Luehe, 


1899. 


Bothriotaenia (part.) 


Ariola, 


1900. 


Abothrium 


Luehe, 


1900. 


Abothrium 


Luehe, 


1910. 



Generic diagnosis: Scolex not exceptionally elongated, with 
two powerful but not especially deep bothria. Segmentation in 
older portions of the strobila mostly insignificant on account of 
superficial wrinkling of the individual proglottides j ripe pro- 
glottides essentially broader than long. Longitudinal nerves the 
lateral borders, dorsal to the cirrus-sac and vagina. Testes ex- 
clusively between the nerve strands. Vitelline follicles of very 
irregular shape, in two broad lateral fields, in part at least be- 
tween the bundles of the longitudinal muscles, the follicles of 
individual proglottides not especially separated from one another. 
Ovary scarcely lobed, more or less bean- or kidney-shaped. Shell- 
gland dorsal to the ovary. Uterus-sac in ripe proglottides an 



undivided cavity, occupying the whole of the medullary parenchyma. 
The openings of the uteri correspond to a more or less prominent 
median longitudinal furrow of the chain of proglottides. 
Type species: A. rug o sum (Batsch). 

Species 1. Abothrium rugosum (Batsch, 1786) . 
1773 Taenia decampollicaris Strussenfelt 1773 : 27 



1781 


Taenia tetragonoceps (part.) Pallas 


1781 




88 


1782 


"Der runzlichter Fischband- 
wurm Tr 


Goeze 


1782 




410 


1786 


Taenia rugosa 


Batsch 


1786 




208 


1788 


Taenia tetragonoceps (part.) 


Schrank 


1788 




46 


1 /90 


Taenia rugosa 


Gmelin 


1790 




3078 


180^ 


Taenia rugosa 


Rudolphi 


1802 




107 


1803 


Rhytis conoceps 


Zeder 


1803 




292 


1810 


Ro th i»io (ienhalus pti^osus 


TudolDhi 


1810 




42 


1816 


Bothriocephalus rugosus 


Lamarck 


1816:168 


1819 


Bothriocephalus rugosus 


Rudolphi 


1819 




137 


1819 


Bothriocephalus rugosus 


Leuckart 


1819 




57 


1845 


Bothriocephalus rugosus 


Dujardin 


1845 




618 


1850 


Dibothrium rugosum 


Diesing 


1850 




591 


1853 


Bothriocephalus rugosus 


Baird 


1853 




88 


1863 


Dibothrium rugosum 


Diesing 


1863 




239 


1867 


Bothriocephalus rugosus 


Olsson 


1867 




53 


1871 


Abothrium gadi 


Beneden 


1871 




56 


1881 


Abothrium gadi 


Monies 


1881 




167 



347 



1839 


Bothriocephalus rugosus 


Linstow 


1889 




2*2 


1889 


Bothriocephalus rugosus 


1'onticelli 


1889 


: 


68 


1890 


Dibothrium ru^osum 


Linton 


1890 


; 


750 


1890 


Abothrium rugosum 


Loennberg 


1890 


: 


22 


1891 


Abothrium rugosum 


Loennberg 


1891 


j 


75 


1892 


Bothriocephalus rugosus 


Matz 


1892 


: 


113 


1894 


Bothriotaenia ru^osa 


Blanchard 


1894 


: 


701 


1896 


Bothriotaenia rugosa 


Ariola 


1896 


: 


280 


1896 


Bothriotaenia ru^osa 


Riggenbach 


1896 


j 


223,228 


1898 


Bothriotaenia rugosa 


Muehling 


1898 


; 


35 


1899 


Abothrium ru^osum 


Luehe 


1899 




39 


1900 


Bothriotaenia rugosa 


Ariola 


1900 




432 


1900 


Abothrium ru^osum 


Luehe 


1900a: 101 


1901 


Dibothrium rugosum 


Linton 


1901 




412,476 


1903 


Bothriotaenia rugosa 


Schneider 


1903a: 


7 


1910 


Abothrium ru^osum 


Luehe 


1910 




26 



Specific diagnosis: IVith the characters of the genus. 
Large cestodes with maximum length, breadth and thickness of 1000, 
7 and 2mm. , respectively. Scolex present only in very young stro- 
bilas, when conical and provided with very weak bothria, changing 
with age to a pseudoscolex of various shapes, usually imbedded in 
pyloric coecum of host. Proglottides at first broad and very 
short, obscured by irregular transverse and longitudinal rugae, 
then gradually lengthening with age until finally they are quadrate 
or longer than broad. 



248 

Cuticula 5^ thick, subcuticula 0.14mm. Small calcareous bodies, 
20j^ in length. Longitudinal muscles in bundles, transverse for- 
ming septa between proglottides. Nerve strands dorsal to cirrus 
and vagina, 45^ in diameter. Two chief excretory vessels anterior- 
ly, passing into 30-35 posteriorly. 

Genital cloaca irregularly alternating, between first and se- 
cond thirds of edges of proglottides. Vagina opens immediately 
behind the cirrus and slightly ventral; no hermaphroditic duct. 

Testes in two lateral fields, discontinuous from proglottis to 
proglottis, ellipsoidal, flattened anteroposteriorly , 40 x 90 x 85 

and 45 to 60 in number. Vas deferens lateral to uterus^sac 
with few coils before entering the cirrus-sac, 350 x 70-80^ . Cir- 
rus-sac ovoid with narrow end outward, 174 to 277^ long by 92 to 
102 in diameter. Cirrus straight in outer half ofi sac, proximally 
coiled or dilated. 

Ovary large, entire, kidney-shaped (isthmus as thick as the 
wings), 0.6mm. wide, occupying the posterior half of the median 
portion of early mature segments. Ova conspicuous, nuclei large, 
10-13|^ in diameter. Oocapt 34^ in diameter. Beginning of oviduct 
Z-shaped. Right and left vitelline ducts join ventrally; common 
duct acts as reservoir. Vitelline follicles entirely within lon- 
gitudinal muscles, discontinuous, intermingling laterally with the 
testes, irregular in shape and size, largest 30, 90 and 70^ in 
length, width and thickness, respectively. Shell-gland compact. 
Uterine duct with only a few coils close to the median line; uter- 













us-sac occupies 


the whole of the 


dorsoventral diameter of the rneduH 


la, very wide and short or irregularly circular or quadrate sur- 


ficially, often lobed, 0.75 to 1.6mm. in transverse diameter, con- 


stantly rounded laterally; openings 


in median zig 


-zag row. 


Eggs, 80 - 


98^ long by 75 - 


92 


wide, shell qui^e transparent. 


Habitat: Intestine df the host with pseudoscolex imbedded in 


a pyloric coecuni 










Host 


Locality 




Collector 


Authority 


Gadus mustela 






Eorke 


Goeze 1782:410 


ft rt 






!7agler 


" " :411 


n aeglifinus 


lYarberg 




Olsson 


uisson iov iio^i 


t? tt 


Eergen 




Loennberg 


Loenn- 
berg 1890:22 


• ff ft 


Grafverna & 
base o 




Olsson 


Olsson 1893:17 


tt ■ 


Arctic Ocean 




Zool.Mus.d. 
i\ .HKau. . n 1 s s • 
Petrograd 


Linstow 1901:281 


n tf 


England 




Nicoll 


Nicoll 1907:71 


" callarias 


Woods Hole, 
Lias s . 




V.N.Edwards 


Linton 1897:431 


rt tt 


Arctic Ocean 




Zool.Mus.d. 

K.Akad.Wiss. 

Petrograd 


Linstow 1901:281 


R TT 


Murman-Kuste 




TT 


" 1903:19 


Tf TT 


Nokujev Id., 
Arctic 




,r (Eaer) 


TT Tf T? 


" lota 


Greiphswald 




Rudolph i 


P.udolphil810:43 













Gadus merluccius 


Rennes ,?rance 


Dujardin 


Dujardin 


1845:617 


" morrhua 


!7arberg 


Olsson 


Olsson 


1867:54 


n ft 


Grand Banks, 
Newfoundland 


Lee 


Linton 


1890:750 


ff IT 


Bergen 


Loennberg 


Loennbergl890:23 


" pollachius 


Rennes 


Dujardin 


Dujardin 


1845:617 




Wart erg 


Olsson 


Olsson 


1867:54 


ff ff 


Bergen 


Loennberg 


Loennbergl890:22 


Tf Tf 


Grafverna & 
Naset, Sweden 


Olsson 


Olseon 


1893:17 


ff Tf 


Millport, Scot- 
land 


Nicoll 


Nicoll 


1910:355 


Lota vulgaris 




Siebold 


Baird 


1853:89 


ff rf 


Memel & Ross- 
itten 


Muehling 


Muehling 1898:35 


ff Tf 


Tvarminne Id., 
Finland 


Schneider 


Schneiderl903b:8 


Morrhua aeglifinus 


England 


Cobbold 


Cobbold 


1858:158 


tf vulgaris 


ff 


Tf 


ff 


" :159 


ff tf 


Belgain coast 


Feneden 


Eeneden 


1871:56 


Merlangus carbonarius England 


Cobbold 


Cobbold 


1858:159 


Merluccius vulgaris 


?farberg 


Olsson 


Olsson 


1867:54 


Me lanogr animus aegli- Woods Hole 
finus Region 




Sumner, 
Osborn 
& Cole 


1913:586 


Microgadustfomcod 


Tf 




Tf 


ff it 


Urophycis tenuis 


TT 




ff 


TT Tf 


Melanograinmus aegli 
finus 


- Passamaquoddy 
Bay, New Bruns. 


Cooper 


Cooper 

(the present pa- 
per) 



351 



Melanogrammus aegli - Eay of Fundy, Cooper Cooper 

f inus Campobello Id. (the prsent pa- 

per) 

■ Tf Freepo/kt,N.S. " " 

Gadus callarias Campobello Id. B 

■ " Woods Hole V.N.Edwards n 



252 

One of the moat striking features of this species la the 
presence of a paeudoaoolex which i3 found embedded in the intes- 
tinal coeca or intestinal wall of the host, from which it is ex- 
tracted only by careful dissection. Goeze (1733 : 412, Figs. 1, 4 
and 5) described a 3colex, somewhat elongated, sagittate and irre- 
gular but otherwise quite comparable to that of other bothrio- 
cephalids, while Rudolphi (1810 : 43, 44) doe 3 not seem to have 
found anything but such a structure in Gaaus lota. Dujardin 
(1845 : 617) wa3 evidently the first tc describe the <: seudo scolex 
by 3aying that, " ... la partie anterieure [of the Strobila] en- 
gagee dans l'appendice pylorique forme une sort de bouchon, un 
cylindre irregulier, cartilagineux, long de 18mm. , large de 4mm., 
ride on toruleux et sans ancune trace d 1 organisation ..." Tr.is 
description, however, was not recognized by Diesing (1850 : 590) 
since he accepted Rudolphi 's diagnosis, namely, "Gaput subsagitta- 
tum, bothriis oblongis lateralibus and Baird (1853 : 8S) evi- 

dently say? two bothria, probably ov/ing to the fact that he was 
dealing with specimens from Lota vulp;aris (vide infra). Cab bold 
(1858 : 158, 159) was well acquainted with the p3eudo3colex, since 
regarding individuals from the Cod ( "Morrhua vulgaris " ) he 3..id 
that "In a Cod examined on the 15th of March, 1885, two specimens 
of Bothriocephalu3 rug03U3 had severally attained a length of 
nearly fifteen inches, and their anterior segments for an inch or 
more downwards, were so firmly impacted within the pancreatic 
coeca, that it was found impossible to dislodge them 7/ithout injur- 
ing the filamentary head and neck. As if to make the anchorage 
doubly 3ure, the cartilaginous thickening of the invaded pancreatic 



353 



coecum had degenerated into a calcareous and contracted cylinder, 
twisted upon itself in various ways." 0l3son (1837 : 54) like- 
wise found a pseudoscolex in this species, which he described as 
being degenerated in Gadus morrhua to a yellow, elongated mass 
viiich disintegrated on contact with water. It wa3 18- 3 5mm. in 
length by about j>wm. in diameter, while its position wa3, as usual, 
in the wall of a pyloric appendage of the host. He al30 figured 
a young strobila from Gadu3 aeglif inu3 , the scolex of - :;hich he con- 
sidered to have been invaginated. Van Beneden (1871 : 56) observed 
the pseudoscolex in his new genus and species, Abothrium gadi , 
which was afterwards considered to be synonymous with A. rugo_sum, 
stating that "113 ont la t£te vers le fond des coecum3 pylori ques, 
percent ordinairement les parci3 et forment, par la gaine, somtent 
dure et entortillee comme une tabulaire, une saillie a la surface 
de cet organe . ■ So far as the writer is aware, he gave the first 
figure of the structure, as it is commonly met with, encased, how- 
ever, by the walls of the pyloric coecum in which it was found 
lodged. Von Lin3tow (1839 : 242) described and figured a scolex 
somewhat similar to that of A. crassum, esoepting that the apex 
was hollowed out to form a six-cornered opening which communicated 
with both bothria. Linton (1890 : 750) found pseudoscolices in 
examples from the codfish, "Gidus morrhua" , which wer much as de- 
scribed by Olsson, since "each of the specimens in this lot ha3 the 
head and anterior part of the body buried in the pyloric caeca, 
•here they have undergone degeneration to such an extent that no 
appearance of bothria remains. Around the parts thus enveloped by 
the caeca is a yellowish waxy deposit, the degenerated tissue of th 



254 



caeca. This adventitious ti33ue invested the worm so clo3ely that 
it "ould be absolutely impossible for the parasite to free itself 
from it3 host." The next important reference to the 3colex was 
by Loennberg (1891 : 75 ) who, while accepting Van Beneden'a new 
genus, Abothrium, referred the species back to rug osum of Batsch, 
and described the metamorphosis of the anterior end of the strobila 
into the well known pseudoscolex, accounting for the various forms, 
3uch as figured here. It is note vorthy , however, that he did not 
3t; ; .te specifically that bothria are present in very young scolice3, 
before thi3 transformation takes place, nor lid he give any figures 
to illustrate the latter. Matz (1332 : 114) described and figured 
a typical 3C0lex for a specimen 36cm. long from Lota vulgaris, 
while Schneider (1903a : 9) in delineating a similar structure for 
the species from the 3ame host, pointed out its great similarity 
to the scolex of B. probosoideus ( = A. cra 3 3um) . Perhaps of sig- 
nificance in connection with the question of the metamorphosis of 
the organ is his statement that "Der ganze Scolex kann 3ich namiich 
durch ver3chiedene Contraction 3einer Muskeln in ein pf eilf oroiiges, 
oder fast cubi3ches, oder sogar sichelf 6rmige3 Gebilde verwandeln." 
Later Johnstone (1907 : 170,171) described the pseudoscolex with 
considerable detail, finding quite the same conditions as did Linton 
On account of never meeting with anything like a typical 3colex in 
adult worms he was led to conclude that "Probably in young codling, 
recently infected, a 3tage of the cestode with such a scolex might 
be found but doubtless with increasing age the changes mentioned 
above occur, and the normal structure of the he=td disappears." And 
la3tly Scott (1909 : 35) made somev;hat similar statements, pointing 



355 

out that " ... no satisfactory description of this part of the worm 
(the pseudosoolex) has yet been published." Th- .3 we 3ee that, 
ap^t from Olsson's (183? : 54) finding in Gadu a aeglif i nus of a 
possible young stage in the degeneration of the scolex of this 
species, no cne has, a3 yet, figured in detail its metamorphosis, 
Loennberg, however, giving the only description of the process. 
On the other hand, a typical 3colex has been described by several 
writers, as pointed out above, for what has been taken to be the 
same species in Lota vulgari 3 , but since there i3 evidence that 
the latter is quite different from the species found in marine 
Gadidae and 3ince the specimens from Lot a maculosa, studied by the 
writer, were found to belong to the well known A. cra 3sum, we must 
attribute a pseudosoolex only to adult3 of A. rugosum, at least 
until the confusion (vide infra) which exi3t3 in the literature re- 
garding the form from Lota can be cleaned up by further investiga- 
tion. 

Two forms of 3colex which were dissected out by the writer 
from the pyloric coeca of Melanogra.ii/flU3 aegli f inus , the Haddock, 
and Gadus cal larias, the Cod, as shown in Fig3. 87 and 88, respec- 
tively, the latter being from the largest specimen at hand, while 
what is doubtless a younger stage in the degeneration of the scolex 
is 3hown in Fig. 86 from the intestine of a Hai:ock, $L series of 
transverse sections of this one, brought out that the internal 
anatomy was quite suggestive of a typical sxolex, that of A. eras sum 
for example. As 3hown in the figure, the structure is somewhat 
flattened; and this flattening wa3 found to be a dorsoventral one. 
While there were only faint suggestions of bothria, especially 



256 

towards the tip, the arrangement of the muscles, nerves and excre- 
tory vessels pointed to this being possibly not far removed from 
the typical form of scolex; and this view is supported by the fact 3 
that it was found free in the anterior part of the intestine of the 
Had.'.ock, altho, unfortunately, the length of the strobila was not 
recorded. Among a lot of material taken from several Haddock two 
examples of the scolex, as it would seem to be at or about the 
time degeneration sets in, were found. The first one, shown in 
Fig. 84, was from the smallest strobila at hand, 23mm. in length, 
while the other, Fig. 85, from an older chain only the anterior 
end of which was present with a length of 33mrn. and maximum breadth 
of 3.5. A3 indicated, the second i3 evidently the older from the 
standpoint of the metamorphosis, since it is more conical and le33 
separated from the neck region which is slightly swollen, while the 
bothria. are disappearing a3 the whole structure is approaching the 
stage represented by Fig. 86. In Fig. 84 are seen somewhat more 
efficient bothria, but the shape of the organ point3 to a consider- 
able amount of degeneration having already taken place. The 3econd 
form of 3Colex is shown in Fig. 87. Here the structure i3 like- 
wise not embedded in the wall of the pyloric caecum in which it is 
found but free in its lumen, the anchorage for the strobila being 
obtained by the close approximation of the mouth of the caecum 
around the narrow neck region and the concomitant swelling of the 
more distal portions. Furthermore, the indications are that a con- 
siderable portion of the anterior end of the strobila is involved 
in the formation of the organ, especially since it is comparatively 
large. The third form observed is shown in Fig. 88, where degener- 



257 

ation has gone on to auoh an extent that there remains only a 
filamentous, horny or cartilaginous yellov/ mas3, deeply imbedded 
in the wall of the caecum, from which it was dissected with con- 
siderable difficulty. Only the tip is shown, there having been 
about 6mm. more to the region where it left the host tissues and 
passed insensibly on to the anterior portion proper of the stroMla, 
■-.ich latter showed only faint transverse wrinkles and no distinct 
division into segments, as is 3een, with some irregularity, however, 
in Fig. 87. This form of pseudoscolex was found, as described by 
Oisson, Linton and Johnstone, to be surrounded by the tissue of 
the coecum degenerated "to a yellow waxy mass which, when freed from 
the surrounding tough tissue, crumbled easily under the dissecting 
instruments. The comparatively great extent of the organ longi- 
tudinally in the wall of the host's pyloric coecum, as the measure- 
ment indicates, need not be emphasized if one has ever endeavored 
to dissect it out entire! 

A3 stated above there is considerable evidence in the 
literature of thi3 species to indicate that the form found in marine 
Gadidae and called A. gadi by van Beneden (1871 : 53) i3 not the 
same as that found in the only fresh-water gadid, viz., Lota . In 
endeavoring to place a number of specimens from Lota macu losa, it 
was found that in many points they agreed with the|de3cription given 
by Matz for A. rugo sum . The 3colice3 are more or less alike, no 
i: seudoscolex (see beloT/, however) being present; the longitudinal 
muscles are not in bundles; the nerve strand3 are dorsal to the 
cirrus and vagina; the genital cloacae are irregularly alternating 
from 3ide to 3ide; the vagina openj ahead of the cirrus instead of 



358 

behind; the testes are continuous from proglottis to proglottis; 
the vitelline follicles are located among the longitudinal muscles 
c-ni are d: scontinuous; and the uteru3-sacs are rounded laterally; 
of whioh characters, however, the position of the nerve 3trands and 
the alternation of the cloacae are applicable to the material from 
the marine Gadidae studied. In many more points, on the other 
hand, the species agreed with A. era 3 sum , 30 that the writer was 
obliged to consider it to belong to that species. A comparison 
of Matz's description with that of Loennberg brings out many differ- 
ences. Loennberg described a pseudoscolex, calcareous bodies, the 
longitudinal mu3cles in bundles, the other sets of pa renchymatous 
muscles as above described, the vagina opening behind the cirrus 
and ventrally, testes discontinuous, vitelline follicles within 
the parenchymatous muscle-sack and also discontinuous, none of which 
characters are to be found in Matz's description, but all of which 
are present in the material at hand from marine Gadidae. It is to 
be noted here that Loennberg accepted the specific name rugo sum of 
Rudolphi instead of the gadi of van Beneden, which as wall be seen 
presently may not be admissable. Going back, then, to the only 
other and the earliest description of the anatomy of the species 
that of Linstow (1389 : 242-5), we meet with similar difficulties 
and confusion. Linstow gave as host 3 for the species, which he 
called rogo sus Hud., Gadus aegl if inus, G. morr ua, Mgrlanjue 
carbo narius, M. po llach ius, Merlucius vul garis, Lota vulgar ij, L. 
molva and Mote 1 1a mustela . Characters in his description not 
applicable to the material studied were: No pseudoscolex, but scolex 
of a rather peculiar shape and structure terminally (vide supra); 



359 

nerve strand 56p. in diameter; 10 excretory vessels anteriorly- 
arranged in two groups of five each; genital cloacae unilateral, 
between the middle and hinder thirds of the edge of the proglottis; 
vagina opening ahead pf the cirrus; length of air r its- sac 0.42mm. 
( ! ) ; ovary 0.14 x 0.13mm.; uterus spherical when obviously young; 
and eggs 59 x 43p/ . Testes with a diameter of 30 y-, vagina 16- 
36f>- . in diameter at the beginning, and two vitelline ducts, be- 
sides a few other minor points in the general anatomy, agree, how- 
ever, with the species as studied by the writer. Thus we 3ee that 
there is by no means anything like complete agreement as regards 
details among the three descriptions by Linstow, Loennberg and 
Matz. But this does not seem to have inconvenienced many of the 
writers since then, notably Ariola (1S00 : 432) and even Luehe 
(1900a) whose references to the position of the vitelline follicles 
and the ventral bow in the vagina are at variance with conditions 
found here; altho Johnstone (190?), Scott (1909) and Niooll (1910) 
were obviously dealing with the form described by Loennberg. 
Schneider (1903a : 7-10) seems to have been the only one who point- 
ed out the differences between the form from Lota and that from 
marine Gadida e . He said: " Bo thr io taenia rugosa gleioht sowohl in 
ihrem Aussehen, als auch in ihrer Anhef tungsweise ausserordentlich 
der Species 3. proboscidea , die in unseren Lachsen ( Salm o sala r ) so 
massenhaft vorkommt. Trotzdem pflegt man aber seit Eudolphi, so- 
viel mir bekannt, immer die in Lota meist vorkommende Form als eine 
getrennte Species aufzufassen unter dem Namen "rugosa" ( Bothrio - 
cephalus ru^0 3us Rud. = Pi oot hr ium rug o sum Die sing u.s.w.), ob- 
gleich die unterscheidenden Merkmale zwischen B. proboscidea und 



260 

B. rugoga , die Riggenbach in seinen "Bemsrkungen ueber das Genu3 
Bothr iotaenia Railliet" ubersi chtli ch zusammenstellt , recht unbe- 
deutend sind und vielleicht loch noch im Rahman der Variationsbreite 
einer einzigen Species untergebracht werden konnen;" and, as regards 
the latter, in a footnote: "Die von If. Luehe ... als Unterscheidungs- 
merkmal vorgeschlagene Lage der Dotterstecke zum Theil (B. rugo sa) , 
bzw. auschliesslich (B. prooos cidea ) zwi3chen den Lang smu skein, 
scheint mir auch nicht genugend constant zu sein, urn als Species- 
inerkmal verwandt werden zu konnen." For material from Lota vul- 
gar is Schneider described a 3colex and segments both similar, a3 
he pointed out, to those of B. proboscidea (= A. crassum ) . The 
arrangement of the genital cloacae, irregularly alternating but 
unilateral for long stretches, the openings of the uteri in a long- 
itudinal furrow, the early form of the uterus- sac and the size of 
the eggs (84.5 x 50-53f*), as described by the same worker all 
agree with A . crassum as studied by the writer (cf. infra). In con- 
clusion Schneider said: "Ueb'rige-ns habe ich, wie gesagt, auch an 
die Examplaren aus dem Museum keine Pseudoscolexbildung bemerkt 
und zweifle daran, dass B. rugo sa und B. gadi ein und dieselbe Art 
sind" and further, "Es ist mir ubrigens bisher noch nicht gelungen, 
B . rugos a oder B. gadi in 0-adus morrhu a de3 Finnischen Ileerbusens 
aufzufinden, obgleich ich zahlreiche Exemplare des Dorsches seciert 
habe, und obgleich B. rugosa in Lota vulgari s hier oft genug vor- 
fcoairat. Auch das scheint gegen die Identitat der Species B. rugosa 
•mit B. gadi zu sprechen." 

Thus we see that there is considerable detailed evidence 
that the species from Lota is not the same as that from the marine 



261 

hosts. We must then go back of Linstow'3 time in order to deter- 
mine, if possible, what is the correct name for the latter. Next 
in retrogressive order is van Beneden's (1871 : 53) description 
of A. gadi , confined to a short footnote which deals With little 
more than the pseudoscolex. So far as it goes this agrees with 
Loennberg'3 A. ragosum and with the material studied by the writer. 
Olsson (1837 : 54) was obviously dealing with the same form which 
he reported from marine hosts only. Die sing (1883 and 1850) copied 
from Rudolphi while Cobhold (1358) had the marine form before him, 
and Baird (1853) had the fresh- water form. In spite of Linstow's 
objection the writer feels certain that Dujardin (184-5) also had 
the 3pecies dealt with here, esx^ecially since his measurements of 
the eggs come nearest to those observed than to those of any other 
writer. It remains then to enquire into Rudolphi 1 s finding and 
description, Leuckart (1819 : 57) copying from him altho at the 
same time remarking that "1st anrnachsten mit den B. proiio scideus 
verwandt, und, wenn er nicht eine Art mit diesem ausmacht zwischen 
B. profro scideusjand B. 3ag ittatus zustellen." For B. rogosus 
Rudolphi ' (1810 : 42-43) described a scolex, quite comparable to 
that of his B. rrofroscidens and to Linstow's description and figure 
of the organ, no neck and segments "primi angusti, fere quadrati, 
insequentes latitudinis raticne habita brevissimi, saepeque inequal- 
es, vel hinc inde angustiores; margines abtusi crassiusculi . " 
This with "Tieque ovaria, neque foramina articulorum vidi ... " and 
the further fact that he obtained his specimens from Gadus iota 
(= Lota vulga ris) , leads the writer to believe that he was not 
dealing with the form present in marine hosts but with a form 



363 



which, if not identical with A. era 3 sum , — his 3. infundi'buli- 
forniis and B. probos ci Jens , — was very close to it. We mu3t then 
go back farther to Batsch (1786 : 208-9) where the species, T. 
rugro sa was named on the basis of Goeze's (1783 : 410) description 
of "Der runzlichter Fi -3 chbandwurm " from Gadus mustela (= Mote 11a 
mustela ) , the marine five-brearded Rockling of Europe, which the 
latter called T. tejtragonoceps Pallas with some doubts, ho7/ever, 
as discussed under the next species dealt with here. Batsch gave 
the following diagnosis of T. rugos a : 

"Taenia (larvata)ci.pite conico cum corpore 
3ubconf luente , papillis lateraliter adnatis usque 
ad apicem capitis, eisque binis: articulis brevi3- 
simis, dilatatis, corpore serrate" 
He used Goeze's Figs. 1-4 and pointed out that he (Goeze) recog- 
nized differences between his specimens and Pallas' T. tetra gono- 
cep-s , for "Er rechnet beyde !~urmer fur eine Art, und die C-lieder 
nebst dem ganzen Korper haben viel Gleichheit, auch die iussere 
Gestalt de3 Kop.fs. Doch sind bey diesem letztern die Saugblasen 
bey weiten nicht so deutlich ^ezeichnet, und stellen vielmehr, wie 
sich Gotze ausdruckt, zwey Backenbarte vor. Die Furche auf dem 
Korper ist auch vorhanden, nur scheint der Eorper mehr gestreckt, 
und am Rande mehr zackig zu seyn." Consequently the correfit name 
of the spcies depends on whether we consider Goeze's description, 
augmented by Batsch' s contributions, to be applicable to the 
material at hand. The largest of Goeze's specimens measured in 
warm water a yard and a half in length by scarcely one-half a line 
in breadth; but the latter is decidedly at variance with his Figs. 



363 

1 and 2 which he said were drawn in "naturlicher Gro33e", in which 
case the width would be from 7 to 15 lines and the scolex about 
12.5 lines (6mm.) in length! For such large specimens, — even 
tho we consider only the first set of measurements, — he described 
and figured nothing of diagnostic value other than a 3colex pro- 
vided with two bothria pretty much of the ordinary type, behind 
this a "distinctly jointed" and "almost cylindrical" neck and along 
both surfaces of the posterior closely crowded segments a median 
longitudinal furrow, all of which characters more nearly agree 
with the probos cideum type of A. era 3 sum (vide infra-) rather than 
with the A. rugosum described above. And since the latter is 
clearly not T. tetragonoceps Pallas as described by Batsch (i.e.: 
304-208), the only course that seems open to the writer is to refer 
the species to van Beneden's Abothrium gadi . However, in view of 
the fact that no material from the European ling ( Lota vulgari s) 
was available for a comparative study, he does not feel justified 
in taking this step, but here retains at least tentatively the 
specific name Abot hrium rugosum (Botseh, 1786) (nec A. rugosu m 
Goeze, 1783). 

The material studied consisted of seven lots from Mel&no- 
grammus aegli finus, and one from C-adus cBularias , from the writer' 3 
Collection; and one lot from the latter host from Coll. Univ, 111. 

















1779 


Species 2. Abothrium crassum (Eloch.1779) 
CFigS. 93-106,) 
Taenia crassa Eloch 1779 


• 


545 




1780 


Taenia salmonis 


LIueller 


1780 


• 


179, 


202 


1781 


Taenia tetragonoceps (part.) 


Pallas 


1781 




87 




1782 


Taenia capite truncato 


Bloch 


1782 




15 




1782 


n Der runzlichter Fischband- 
wurm" 


Goeze 


1782 




410 




1782 


Taenia r>r»oboscis snilla 


U V V fcl V 


1782 


• 


417 




1786 


Taenia totragonoceps 


Ratsch 

XJ W kJ \s XX 


1786 


• 


204 




1786 


Taenia ppoboscidea 


P&tsch 

A ' Vw VIA 


1786 


• 


212 




1790 


TplPfll?? R A "I mOTl 1 ^ 


("J-Tnp 1 1 n 

VjTXXlw X X XX 


1790 

X / <J\J 


• 


3080 


1790 


Tapnia salvplini 

ICO VlilU uuXVvJ.XilJ. 


r>ph t»p nlr 

u o n i Uj liiV 


1790 


• 


125 




1793 


Tfl pn ia 1 vp 1 l n i 


QpTi t>o n If 

OWXI alXxi. 


1793 


• 


141 




1795 


Taenia salmonis 


Rudolphi 


1795 


• 


17 




1802 


TflPnifl q a 1 mrm 1 <? 
X ct t/ ii X ol o <X J-IUU 11 X o 


X'UOO 


180? 


• 


308 




1802 


X CI. i X X CX \J 1 ^UUOOXUvd 


HUUU X^JXX X 


1802 


• 


106 




1803 


Rhytis salvelini 


Zeder 


1803 


• 


292 




lolU 


Ro t.h y»i Of. pnh a 1 n "npohnqr* i r)pa 

XJU Ulll X vuw|JJlclXUO pi UUVOuXUCa 


Pudnl nh i 

i i. LIU. V X Ull X 


1810 




39 




1810 


Rrtf Vcpi hp . i nf nnr) t 1 i "Fn y»ttii q 
lu u.u x'Ju t xnx unuxuuxxxui uxo 


Pnnn I -nit i 

l-UU.U X UXX X 


lftl 

lUXW 


• 


46 




1816 


Bothrioc. proboscidea 


Lamarck 


1816 




582 




|oiy 


Bothrioc. proboscidea 


Rudolphi 


1819 




137, 


472 


1819 


Bothrioc. infundibulif ormis 


Rudolphi 


1819 


• 


137, 


473 


1819 


Bothrioc. proboscidea 


Leuckart 


1819 




38 




1819 


Bothrioc. infundibulif ormis 


Leuckart 


1819 


: 


42 





285 



1843 


Bothrioc . 


salmonis umblae 


Koelliker 


1843 




91 


1844 


Eothrioc. 


proboscideus 


Bellingham 


1844 




252 


1844 


Bothrioc . 


infundibulif ormis 


Belliggham 


1844 




253 


1845 


Bothrioc . 


proboscideus 


Dujardin 


1845 


: 


615 


1845 


Bothrioc . 


infundibulif ormis 


Dujardin 


1845 




616 


1846 


"Bothriocephalus du Saumon" 


Blanchard 


1847 




116 


1850 


Dibothrium proboscideum 


Diesing 


1850 


: 590 


1850 


Dibothrium infundibuli forme 


Diesing 


1850 




590 


1853 


Eothrioc. 


proboscideus 


Baird 


1853 




88 


1853 


Bothrioc. 


infundibulif ormis 


Baird 


1853 


: 


88 


1863 


Dibothrium proboscideum 


Diesing 


1863 


: 


242 


1863 


Dibothrium infundibulif orme 


Diesing 


1863 


: 


242 


1867 


Bothrioc. 


proboscideus 


Olsson 


100/ 




53 


1871 


Bothrioc. 


proboscideus 


Beneden 


i-O / X 




69 


1878 


Bothrioc. 


infundibulif ormis 


Linstow 


lO * o 




263 


1884 


Bothrioc. 


infundibulif ormis 


Zschokke 


1884 




21 


1889 


Bothrioc. 


suecicus 


Loennberg 


1889 




35 


1892 


Bothrioc . 


infundibulif ormis 


Matz 


1892 




110 


1893 


Bothrioc. 


infundibulif ormis 


Olsson 


1893 




17 


1893 


Bothrioc. 


proboscideus 


Olsson 


1893 




17 


1894 


Bothriotaenia infundibuli- 
f ormis 


Blanchard 


1894 




701 


1896 


Bothriot. 


infundibulif ormis 


Ariola 


1896 




280 


1896 


Bothriot. 


infundibulif ormis 


Riggenbach 


1896 




223 


1899 


Abothrium 


crassum 


Luehe 


1899 




39 



266 



1900 


Bothriot . 


proboscidea 


Ariola 


1900 : 


433 


1900 


Abothrium 


crassum 


Luehe 


1900a: 


97 


1909 


Bothrioc . 


proboscicieus 


Scott 


1909 : 


78 


1910 


Abothrium 


crassum 


Luehe 


1910 : 


26 


1910 


Abothrium 


crassum 


Ward 


1910 : 


1184 



Specific diagnosis: With the characters of the genus. Large 
cestodes with maximum length, breadth and thickness of 870, 6 and 
2mm. , respectively . Scolex variously shaped; usually rounded pos- 
teriorly and truncated anteriorly; with prominent bothria and ter- 
minal disc. First segment may or may not be elo^ated to form a 
short neck. Proglottides at first broad and short or more Qua- 
drate, cuneate or infundibulif orm in shape; then, in the middle of 
the strobila, five or more times broader than long; and finally 
posteriorly quadrate or as long as broad. Usually a median lon- 
gitudinal groove down each surface of the strobila formed by emar- 
ginations on the posterior borders of the segments. 

Cuticula 4 to 5 fiA- thick, subcuticula60 to 100 y~ . Calcareous 
bodies (?) absent in adult strobilas. Longitudinal muscles not 
in bundles; no muscular septa between proglottides. Nerve strands 
40^ in diameter, dorsal to inner end of cirrus-sac. 12 chief ex- 
cretory vessels, 6 on each surface just within the transverse 
muscles, reduced to 6 or 8 anteriorly. 

Genital cloaca irregularly alternating, but unilateral for 
long stretches; from one-third to one-half way along the margin 



367 

of the proglottis. Vagina opens ahead of and slightly ventral to 
the cirrus; no distinct hermaphroditic duct. 

Testes within the nerve strands, pseudostratif ied, continuous 
from joint to joint; elongated dorsoventrally , 95-115 x 70-100 ja- ; 
40 to 150 in number. Vas deferens lateral, elongated, with few 
coils before entering the cirrus-sac, 350-600 x 150-180 ^ in dimen- 
sions. Cirrus-sac ovoid with narrow end outward, 130-380 x 60-150|jl 
Cirrus proper an almost straight tube in outer half of sac. 

Ovary comparatively small, irregular or somewhat lobed, with 
thick isthmus, o.8mm. wide by 0.13 long. Oocapt 40)^ in diameter. 
Usually two ventral vitelline ducts unite to form a common duct 
which does not act as^reservoir . Vitelline follicles irregular in 
shape and size, among the longitudinal muscles or outside of them, 
discontinuous. Shell-gland small, compact, dorsal. Uterine duct 
with on}y a few coils near the median line. Uterus-sac transversely 
elliptical or quadrate and somewhat lobed, rounded laterally, fil- 
ling up almost the entire proglottis when gravid; opening in the 
median line opposite emarginations of segments. 

Eggs, 45-115 x 30-75j^, 6void or ellipsoid in shape. 

Habitat: In th^; pyloric coeca and intestine of the host. 

Host Locality Collector Authority 

Salmo salar - — Borke Goeze 1782:417 

" " Batsch Stiles and 

Hassall 1912:395 

■ ■ Rudolphi 













268 


Salmo 


salar 




Zeder 


Stiles & 
hassall 




n 


tt 




r.uu.oxpn i 


T a1 t\T^ t 

j uQoipm 


1 JR1 Pi. AD 


rr 


tt 


uTypnswaia 


ft 


tr 




ff 


n 


T wrt 1 ft in ri 


Poll i yy '~"V» o vn 


Bellinghaml844:253 


ft 


tt 


Paris 


Tin t v» 

l»uj arciin 


Dujardin 


1845:615 


Tf 


n 




M . U • V . 


Diesing 


1850:590 


ft 


tf 




Giebold & 
Johnston 
(Coll. Frit. 

MUS • } 


Baird 


1853:88 


tf 


tt 


\7arberg 


Diss on 


Olsson 


1867:53 


ff 


ft 


Belgian coast 


Beneden 


Beneden 


1871:69 


ff 


tr 


'.Yarnemunde 


r> _ _ v 1,1--. 

ZiScnoKice 


Braun 


1891:55 


n 


tf 


Uaset 


01s son 


Olsson 


1893:17 


ft 


tt 


rcnine k . ,i:asei 


Z schokke 


Zschokke 


1896:776 


S almo 
bill 


salar no- 

LS 


Murman-Kuste 


Zool.Mus. 
Kais.Akad. 
T . v iss . ,Petro 
grad 


Linstow 


1903:20 


Salmo 


salar se- 

) 


Lalce Sebago,Me. 


Wafrd 


Ward 


1910:11 
(84 


baimo 


ajbpinus 




Mus . Vienn. 


Rudolphi 


1819:137 


tf 


n 


J emtland 


Olss on 


Olsson 


1876:149 


rt 


tt 


Lakes ITackten, 
Stors jon,Lock- 
nes jon 


tr 


tt 


1893:17 


tt 


carp 10 


L.traraa , i t.aiy 


Nmni 


Stossich 


1890:7 


tt 


caspius 


Karabugas -S trass e 


Maximo vie 


Linstow 


1903:20 


tt 


f ario 


Ireland 


Bellingham 


Bellinghaml844:252 













359 


Saline 


f ario 


Rome 


Condorell i 


Ariola 


1900:435 


tr 


TT 


Vyg-Fluss 


Danile vski j 


Lins tow 


i orio . on 
iy(Jo: 


n 


hucho 




Mus.Vienn.& 
Eremser 


Rudolph i 


1819:472 


rt 


Tf 




Diesing 


diesing 


1 D CO . con 


tf 


lacustris 


Benacd 


Largaiolli 


Ariola 


1 AAA ./IOC 

lyUU: 435 


tt 


Hamaycush 


Shoal Id., Lake 
Superior 


Milner 


TT 


TT If 


tt 


salvelinus 




Schrank 


Stiles & 
Hassali 


i m O • A AO 


fT 


TT 




Zeder 


TT 


:oUd 


rr 


TT 




Mus • Vienn. 


r^UQOipnl 


lOlQ, 1 OO 


TT 


TT 




Diesing 


Diesing 


l ocn. col 

looU: oyi 


TT 


siecowet 


Outer Id., Lake 
Superior 


Milner 


Ariola 


1900:435 


9 n 


thymallus 




uoii • v ienn . 


Leuckart 


1D1Q, ^O 


TT 




M • C • V • 


Diesing 


looU: oyi 


TT 


trutta 


Ireland 


Eellingham 


Eellinghaml844:253 


TT 


TT 




Coll.Brit. 

IV:US • 


Baird 


1853:88 


Tf 


Tf 


Lakes Storsjon, Olsson 
Halen,Refunds- 
s jbn,Salls jon,& 
Ockes j on, Jemt land 


Olsson 


1893:17 


TT 


TT 


Murman-Kiiste 


Zool.Mus .d. 
Kais.Akad. 
Tiss. ,Petro- 

grad 


Linstow 


1910:281 


TT 


umbla 




M.C.V. 


Diesing 


1850:591 


Tf 


TT 




Zschokke 


Zschokke 


1884:21 











270 


Core^onus fera 




r ' c? /■» V\ r\ Tr Ir r\ 

u s cnoKKe 


Zschokke 


1 QQ/1 . 91 

loo4: cil. 


lavaretus 


Lakes Storsjtfn, 
and iiaicien 


Olsson 


Olsson 


1893:17 


" oxyrhynchus 
maraena 


IVarberg 


Olsson 


Olsson 


1867:53 


lrutta iano 


Genf ersee , Easel 


Zschokke 


Zschokke 


loyb: fib 


" lacustris 


ruiine rt.,.basei 


tt 


n 


tt tt 


tt tr 


Bodensee 


hoi er 


xioi er 


iyu4: &<oi 


" salar 


East Prussia 


lauenimg 


Muehling 


1 O QQ . Q C 

loyo : o o 


Lrutta 


W ar n e nun d e 


Zschokke 


Braun 


1 ftQI . £C 

loyi : oo 


rf var i a"b i 1 i s 




tt 


ZscnoKKe 


1 QO/I .91 


Tti vrfic? line ttii 1 cto t» t c 

lllj iliO- JL -L U.O VtlJ-^cXI ifi 




tt 


rt 


rf tt 


If n 


Stbrs j6n,Jemt- 
land 


Olsson 


Olsson 


1893:17 


ft ft 


Eaikal-See 


Zool.Mus.d. 
Kais.Akad. 
Wiss. ,Petro- 
grad 


Linstow 


1903:20 


Esox lucius 




Zschokke 


rr _ „v -1,1--, 

ZiScnOKKe 


1 QfM . 9 1 


rerca i luvia tins 




tt 


tt 


rr tt 


Osmerus eperlanus 


B6nan,Gestri- 
cia,Gulf of 
Bothnia 


Olsson 


Olsson 


1893:17 


Clupea harengus 


Os t se e 


Schne ide r 


Schimider 


i on? . 9R 
i y u<o : oo 


Lota vulgaris 


Stbrs 3 on, Jemt- 
land 


Olsson 


Olsson 


1893:17 


rf tt 


Bvina-Fluss 


Danile vski j 


Lins tow 


1 QfiQ . 9H 

lyuo : (~>\j 


irout 


boon lay 


.. l ii lamson 


OCOX.t 


1 QflQ. 7ft 

iyuy : i o 


Cris tivomer namay** 


Slant's Tomb 
Id. , Georgian 

Bay, L.Huron 


Cooper 


hooper 

(the present paper) 







271 



Cristivomer nama.^ 
cush 



Port 6redit,0nt., Cooper 
Lake Ontario 



Cooper 

(the present gaper) 



" Lota lota " 
Lftta maculosa 



L.Temagami,Ont. 
Charlevoix, Mich . 
Pentwater,Mich. 
Charlevoix 
Port Credit 



II. B.Ford 



Cooper 



Coregonus clupei - Giant's Tom b 
formis Island 



272 

Thi3 species, originally given the specific name of 
Taenia crassa by Bloch (1779 : 545), was on the one hand confused 
With A. rugo s um and on the other given the new name Taenia probo- 
scis suiil a by Goeze (1782 : 410-11, 417, re3p.) according as it 
me found in Gadus or in Salm o salar , which confusion was evidently 
due to the fact that the latter followed Palla3 (1731) in calling 
it T_. tetra ganoceps . In a footnote he considered, in fact, that 
T. tetr agonoceps Pallas, Taenia crassa Blech and T. capite tjanca to 
Blach were all synonymous. But in spite of this he expressed 
doubt on the synonymy of the forms from the Gadidae and from the 
Salmonidae. As the above synonymy indicates the 3pecies was then 
known for a number of years under at legist two names, Botnri oce- 
cephalu3 i n f und i bu 1 i f o r m i s and B. probosoidelg which were used by 
Rudolphi (1810 : 46, 39) for the two forms from freshwater and 
marine salmonids in general. It was not until 1S84 that Zschokke 
made a detailed cornp; risen, — it is true of external characters 
mostly, — of the two species from a number of different hosts, and 
showed that they must be considered only different forms of the 
same species. Later investigations into the anatomy by Matz (1892 : 
110), who, however, studied only the pro bosc ideu3 form from Trutta 
trutta and Salmo salar , have been considered to have established 
this contention, altho Olsson (1893 : 17) still reported both of 
the older species with some doubt a3 to the use of the name B. in- 
fundibu liformis . BlftHohard (1894 : 701), Ariola (1896 : 230) and 
Riggenbach (1895 : .223) evidently accept©<i.only the latter specific 
name, while Luehe (1899 : 39) first used the combination Abothrium 
era s su m _(_Bloch) which is now generally accepted. Ariola (1300 : 



273 

433), it might be mentioned finally, called the species ( Bothrio- 
taenia prob oscidea. (Batsch), thus disregarding the fact that Batjjsh 
(1786 : 212) renamed Bloch's T. crassa . 

As regards the scolex, various forms of which are shown 
in the figures, the s: ecimens from Lota maculos a, the common 
Ling or Burbot, need special mention, since, a3 shown in Fig. 38, 
the terminal disc and anterior half of the or^an is in many instanc- 
es greatly swollen to form a sort of pseudoscolex which is perhaps 
veil adapted to maintaining its position in the narrow pyloric 
caecum of the host. But this modification was found only in the 
older strobilas of the four lots examined. In the younger chains 
the scolex is as shown in Figs. 96 and 97 which are drawn to the 
same scale as that of Fig. 38. The largest with this first form 
of scolex was 22mm. in length by 1.5 in breadth while the shortest 
with the swollen end was 45 x 1.5mm., so that somewhere between 

OS 

the lengths of 20 and 45mm. the metamorph^s takes place in all pro- 
bability. No distinct intermediate stages were seen altho the 
smaller scolices were varied in shape and degree of intactness. 
The latter might seem to point to the condition being due to me- 
chanical or physical means, but this is offset by the fact that 
the material was found to be in good histological condition when 
sectioned; hence the idea of a possible metamorphosis. Unfortu- 
nately the material did not permit of anything more to be said on 
this matter. 

In the anterior segments of considerably relaxed or espe- 
cially young strobilas something of the manner of segmentation can 
be seen. This was found to take place much as in the genus 



274 

Both rioc ephalic , altho the writer was not able to distinguish the 
primary segments to hi 3 satisfaction. IThat was considers! to be 
such is shown in Fig. 99, a stretch of segments beginning 27mm. 
from the anterior end of the strobila in question. The idea of 
dominence of the anterior portions over the posterior portions, 
as dealt with under B. scorpi i , is here brought out very nicely. 
In the proboscideus type of strobila the same method of subdivision 
was followed in the anterior segments, altho with greater difficul- 
ty on account of the fact that the segments are so closely crowded 
together longitudinally. 01s3on (1S57 : 53) noticed the subdivision 
of the segments producing an alternation of larger and smaller ones, 
but he considered it to be an arti c ulat la spuri a similar to that 
described by Yfagener (1854 : 69) for Anphio otyle heterople urum and 
by Krabbe (1365 : 37) for B. soorjg dj and other species. Later 
Olsson (1393 : 17) stated that transverse divisions occurred in 
B. infund ibulif o rmis a 3 well as in B. prob oscideus . 

The youngest lots of material, studied by the writer were 
two taken from Lota maculosa from Lake Ontario, off Port Credit, 
Ontario, and one from the intestine of a young Cristivomer 
namaycush from the same locality. The lot from the Lake Front 
contained all stages from that shown in Figs. 104 to the largest 
which by comparison with adult specimens from the same host were 
found to belong to this species. While no stages were found be- 
tween that shown in Fig. 104 and that in Fig. 102, — altho two 
others were only slightly larger than the latter, — it seems rea- 
sonable to consider the latter itself to belong to this series and 
to represent the earliest stage of the same. Figs. 105 and 106, 



375 

of two later stages, are given to show the manner of beginning of 
the segmentation and the early drop/ing off of two or more very 
immature segments from the posterior end. The first indications 
of this is probably represented in Fig. 104, altho the strooila in 
Fig. 105 does not 3how it. The relative ages, howeve:. , of these 
two is difficult to state definitely since the first one is more 
contracted longitudinally than the other. On the other hand, two, 
intermediate in length between those shov/n in Figs. 105 and 106, 
were indented posteriorly, thvs showing that some of the earliest 
segments had already been lost. Thus we see that at a very early 
period in the development of the 3trobila of this species a few of 
the first formed segments are lost in much the same way as the 
bladder of the cy3ticercus of the taenioid cestodes is cast off in 
the final host. 

The material studied consisted of five lots from Salmo 
salar , eleven from Criati vomer namaycush, two from Cofcgonus clupei- 
formis , and two from Lota maculosa , all from the writers collec- 
tion; and, eight lots from C*. namaycush , and two from L. maculosa 

10TV 

from the Collect-^ of the University of Illinois. 



276 



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te Haarlem, 2 Verz. Deel 13, 112pp., pls.l-36A. 



397 

Ward, H. B. 

1910. Internal Parasites of the Sebago Salmon. Bull. Bur. 
Fisheries, 28, 1908 : 1153-1194, lpl., lOfigs. 
We inland, D. F. 

1858. An Essay on the Tapeworms of Man, etc. x 93, 12 figs. 
8 . Camb.(Mass.) 

1859. (Re Schistocephalus in Bhynchichthys gronovii ). 
13 Jahresber. d. Ohio-Staats-Landbaubehbrde , 1859 :566. 
(Affcfcr Leuckart.) 

Willemoes-Suhm, R. 

1869. Helminthologisches Notizen 1.1. Zur Entwickelung von 
Schistocephalus dimorphus Creplin. Zeit. f . Wiss. Zool., 
19 : 470-472. 

1870. Ibid II. Idem 20 :94-98, Taf . X. 
Wolf, Eugen 

1906. Beitr'age zur Entwickelungsgeschichte von Cyathocepha - 
lus truncatus Pallas. Zool. Anz., 30(1-2) :37-45, figs. 
1-5. 
Wolffhugel, K. 

1900. Beitrag zur Kenntnis der Vogelhelminthen. Diss. 

(Basel), 204pp., 7pls., 114figs. 4 . Freiburg in Br. 

Zeder, J. G. H. 
1800 

^Erster Nachtrag zur Naturgeschichte der Eingeweidewiirmer, mit 
zufassen und Anmerkungen herausgegeben. xx 320pp., 6pls., 
4 . Leipzig. 



398 

Zeder, J. G. H. 

1003. Anleitung zur ITaturgeschichte der Eingeweidewiiriner . 
xvi 432pp., 4pls., 8 . Bamberg. 
Zernecke, E. 

1895. Untersuchungen uber den feineren Fau der Cestoden. 
Zool. Jahrb., Anat., 9(1), 25 :92-161, pis. 8-15. 

Zschokke, F. 

1884. Recherches sur 1* organisation et la distribution zoo- 
logique des vers parasites des poissons d'eau douce. Arch, 
de Biologie, 5 : 154-243. 

1896. Zur Faunistik der parasitischen Wurmer von Siisswasser- 
fischen. Centrbl. Bakt. u. Parasit., 19(21) : 772-784; (21): 
815-825. 

1903. Die Arktischen Cestoden. Fauna Ark. Ill sl-31, 2pls. 
1903a. Marine Schmarotzer in Giisswasserf ischen. Verhandl. d. 
naturf. Gesellsch. in Basel, 16 :118-157, 2pls., 23figs. 



EXPLANATION OF PLATE I. 

Fig. 1. Ligula intestinalis , scolex of larva. 

Fig. 2. , anterior end of adult. 

Fig. 3. , median portion of a transection 

thru the genital cloaca. 

Fig. 4. Ligula intestinalis , union of vagina, oviduct and vitel- 
line duct. 

Fig. 6. Ligula intestinalis , toto of larva from Micropterus dolo - 
mieu. 

Fig. 7. Schistocephalus sAlidus , anterior end of larva. 
Fig. 8. w , median portion of transection 

thru the ovary. 

Fig. 9. Schistocephalus solidus , median portion of transection 
thru the seminal vesicle and cirrus-sac. 



EXPLANATION OF PLATE II. 
Fig. 5. Ligula intestinalis , larva from liver of Gasterosteus bi- 
spinosus . 

Fig. 10. Haplobothrium globulif orme t outline of smallest larva at 
hand. 



Fig. 11. Haplobothrium globulif orme , scolex, toto. 

Fig. 12. n " , primary strobila, toto. 

Fig. 13. " " , secondary scolex, surficial 
view. 

Fig. 14. Haplobothrium glabulif orme , same, lateral view. 

Fig. 15. , transection thru scolex. 

Fig. 16. , the ganglionic mass. 

Fig. 17. w , transection thru proboscis 



bulb. 

Fig. 18. Cyathocephalus americanus , scolex, toto. 



EXPLANATION OF PLATE III. 
Fig. 19. Cyathocephalus americanus t transection thru ovarian is- 
thmus . 

Fig. 20. Cyathocephalus americanus . diagram of medial sagittal 
section. 

Fig. 21. Cyathocephalus americanus , frontal section of ripe pro- 
glottis. 



Fig. 


22. 


Cyathocephalus 


americanus, oocapt containing an ovum. 


Fig. 


23. 


Marsipometra hastata 


i 


scolex, surficial view. 


Fig. 


24. 


n 


n 


9 


same, lateral view. 


Fig. 


25. 


TT 


TT 


9 


transection thru ovarian isthmus. 


Fig. 


26. 


TT 


T 


t 


toto of ripe proglottis. 


Fig. 


27. 


IT 


TT 


1 


genital cloaca from frontal section 


Fig. 


28. 


TT 


TT 


> 


cirrus-sac from a transection. 


Fig. 


29. 


TT 


TT 


f 


larva. 


Fig. 


30. 


TT 


TT 


f 


older larva. 







l?YPT.AT T ATTni\T OF PT.ATF TV 

ljAi JjfU.il 1 Iwri VJr Xlmllj IV. 


r lg. 


91 

ox • 


lriacilU pilU I Ub | ldl V cl j I U JUb tub It j pr j o UI X X C X d.X VI sw . 


r lg . 


OC/ * 


^ bet lilt; j Xdl»t<IclX VXcVV. 


TP i a- 
r lg. 


99 


coma rtnci A "p *f Vt p 4r»irlov"t4o at* 1*i a aIt* c? 

j sdiiioj one ux uric uiiuciii/b ox iiooks • 


r ig. 


oft . 


*? "1 r> v Q nnrlnl neiie 4" tttv o cnr»"Pir>TCiT tttotit 
{ XcLX Vo. j IlULlUIUoUb pc j b UX 1 lliaJ. V ±CW • 


r lg • 


9R 


y b CtillfcJ XCl.l/CXCLX VXtfW. 


r lg. 


9fi 


COTTD o T T 1 1 ^ Id >n T ayi ^ tt n nm 

^ bo.lL" f X 'a lL.cil b j cnu V 1 tw • 


r lg • 




y od.li±tjj bUXIXCXexX VlCW • 


Fi c . 
r x& . 


9ft. 


Ro i" Vi t»i np p"rVh p, 1 n rpottiit ^poIpy ^nypi p, 1 vi pw. 


r lg • 


9Q 


j bdliltJ ^ XclLCXclX VlcWi 






tKyipo prttPTHO'P n'Pi'mflT'V p crmPTi f <3 . 


r lg. 


ftX . 


Aria ~P O 4" Q V> l^Q n 1 pri TATA 

f UIlc Xdl bXlcX DdtK.j dioU ID tO • 


r xg, . 


4-2. 


y emu wici oil u »» x j.^ i i uuui i> x v c 






T>nn l TnonT e TATA 
I UUXIIlcIl li b j tOOU. 


cr 

r lg • 




D U bXl I lUtcpildlUo oOUX fJ X X 9 1>I ctll bcO l» X U Ii l»il 1 U UVo-ij » 


ErXg • 


AA 


Y1 A T> 4" 1 A VI A -p etTJAVillci cTn AW 1 ncr P Y • 
j JJvJ.1 OXUll UX b li UUllc. oXlUVYXil^ CA" 




cretory vessels. 


Fig. 


45. 


Bothriocephalus scorpii, outline of mature segments. 







VYPT ANA TTfYM (W PT A TV V 


e lg. 


AC. 
4tO • 


D Uiiri pilal ub bCOrpil ( OOLO Oj. EdLUrc prO£i, 1 X» LI Q S • 


Pin- 

r lg* 


4.7 


y iiitju.id.ri Do^i i) i;ai Sccuj-Oii coinpo - 






site • 


r ig. 


Aft 


DU J. V t? pil cx lub bOUrpXJ, | kU bU Ux LnU o o^iufc; 11 l> b • 


r xg • 




Ti o ti t l An rx'P cpr< t i nn c Vi rmr i n -/ n r> _ 
y yui UXUI1 UX oct 01UI1 bllUWXIl^ Ull" 






ion of vagina with oviduct* 


Fig 50. 








view • 


Fig* 


51. 


D. CldvlCepS , baluc j Id. Udell VlcW. 


Fig* 


52, 


9 X X UiU IAXJ-J.CX 1 UD yl clUClj O wulCA j CIXIXXUXCX 






view* 


Fig. 


53. 


D. CxclVlOcpb , IrOIIl iliUp. £4 X 1) UUbUb , UI cilibc C OxOil i»iiru uvciry • 


Fig. 


54. 


oomo 4 f\ 4 r*\ a-P no ftivio r\Y»/"*rrli^4 4T/^ao 

f b&nic j to to ox nid ourt? prugio LLiiiob • 


Fig. 


55. 


, same, median sagittal section. 



EXPLANATION OF PLATE VI. 

Fig. 56. B. cuspidatus . scolex, surficial view. 

Fig. 57. n " , same, lateral view. 

Fig. 58, " , transection thru an anterior segment. 

Fig. 59. " , transection thru ovary of mature proglottis 

Fig. 60. n , to to of ripe proglottides, posterior in 

deeper optical section. 

Fig. 61. B. cuspidatus , median sagittal section , composite. 

Fig. 62. tr , young egg, showing an early stage in de- 

velopment. 

Fig. 63, B. cuspidatus , ol^der egg, many celled stage. 



EXPLANATION OF PLATE VII. 

Fig. 64. B. manubriformis , scolex, surficial view. 

Fig. 65. 9 w , same, lateral view. 

Fig. 66. " , anterior primary segment. 

Fig. 67. 9 , transection thru anterior regiom. 

Fig. 68. " , toto of mature proglottides. 

Fig. 69. n , cirrus-sac and vaginal bulb in tran- 

section. 

Fig. 70. B. manubrif ornis , transection thru uterus opening. 

Fig. 71. B. occidentalis t scolex, affc&r Linton. 

Fig. 72. 9 , cirrus-sac in a transection. 

Fig. 73. Clestobothrium crassiceps , scolex, surficial view. 
Fig. 74. , same, lateral view. 

Fig. 75. , same, terminal view. 

Fig. 76. , toto of scolex and anterior end 



EXPLANATION OF PLATE VIII. 
Fig. 77. C. crassiceps , toto of youn- strobila. 
Fig. 78. " , primary segment with reproductive rudi- 

ments, toto. 

Fig. 79. C. crassiceps , segments showing spurious articulations, 

Fig. 80. " , transection thru ovary. 

Fig. 81. " , toto of mature proglottis. 

Fig. 82. n , median sagittal section, composite. 

Fig. 83. " , four consecutive sections thru union of 

vagina and oviduct, showing the receptaculum seminis. 
Fig. 84. Abothrium rug o sum , scolex of young strobila. 
Fig. 85. w , later stage in degeneration of same, 

Fig. 86. n " , still later stage. 



EXPLANATION OF PLATE IX. 

Fig. 87. Abothrium rugosum, pseudoscolex from lumen of pyloric 
coecum of host. 

Fig. 88. Abothrium rug o sum , pseudoscolex imbedded in wall of coe 
cum. 

Fig. 89. Abothrium rugosum, transection thru ovary. 

Fig. 90. " , terminal excretory vesicle. 

Fig. 91. , frontal section of mature proglottis 

Fig. 92. " , union of vagina and common vitelline 

duct with oviduct. 

Fig. 93. Abothrium crassum , scolex of specimen from Salmo salar, 

surficial view. 

Fig. 94. Abothrium crassum , same, lateral view. 

Fig. 95. n , scolex from Cristivomer namaycush . 

Fig. 96. " , scolex from Lota maculosa . 

Fig. 97. " ? same, lateral view. 



EXPLANATION OF PLATE X. 
Fig. 98. Abothrium crassum , enlarged scolex from Lota maculosa . 
Fig. 99. " n , anterior segments of strobila from 

Coregonus clupeif ormis , toto. 
Fig. 100. Abothrium crassum , transection thru ovary of specimen 

from Salmo salar . 
Fig. 101. Abothrium bras sum , toto of ripe proglottis of chain 

from C. clupeif ormis . 
Fig. 102. Abothrium crassum, plerocercoid from C. namaycush t sur- 

ficial view. 

Fig. 103. Abothrium crassum , same lateral view. 
Fig. 104. " , young strobila from same host. 

Fig. 105. ,T , older strobila from same host. 

Fig. 106. " , still older strobila, showing drop- 

ping off of segments posteriorly. 




3 



9 



PLATE I. 




PLATE II. 




/8 



5A 



PLATE II. 




PLATE III 



Z 1 



3o 



>z?> 



PLATE III. 




PLATE IV. 



31 ^ 



3V 



3/ 3 



3* 



3? 



4? 



^3 



1* 



PLATE IV. 

I 




PLATE V. 



6? 



fT3 



57) 



fx. 



*7 



(St 



t 



PLATE V. 




PLATE VI 



7 



Ci 



Co 



5? 



5* 



PLATE VI. 




PLATE VII. 



to 



7i 




7r 
it 

(J 



PLATE VII. 




PLATE VIII. 



S7 



6V 



7? 



*9- tfs' 



PLATE VIII. 




PLATE IX. 

n 

n 



n 



PLATE IX. 




PLATE X 



W3 W 



Ml 

/or 



PLATE X 








' VITA. 






Arthur Reuben Cooper. 


1888 




Born at Brinston, Ontario, Canada, October 30. 


1896- 


•1901... 


Attended the local public school. 


1901- 


■1906. . . 


Ait T "J T • TT • 1 At 1 It • r\ i * 

Attended Iroquois High School, Iroquois, Ontario. 


1906« 




Entered the University of Toronto with the Seventh 






Edward Blake Scholarship in General Proficiency at 






the Honor Matriculation Examinations. 


1906- 


■1910... 


Pursued the honor course in Biology in Victoria Coll- 






ege, obtaining the Gold Medal in "The Science of Bio- 






logy" . 


1910. 




Obtained the degree of Bachelor of Arts from Victoria 






College, University of Toronto. 


1909- 


■1911... 


Class assistant in general Biology. 


1911. 




Obtained the degree of Master of Arts with "Honors" 






from University College, University of Toronto. 


1911- 


■1912... 


Demonstrator in zoology, University College. 


1909- 


-1912... 


Four summers at the Lake Biological Station on Geor- 






gian Bay, Go-Home Bay, Ontario. 


1912- 


■1914... 


Fellow in Zoology, University of Toronto. 


1913- 


■1914... 


Two summers at the Marine Biological Station at Saint 






Andrews, New Brunswick. 


1914- 


■1915... 


Demonstrator in Zoology ; University of Toronto. 


1915- 


■1916... 


Fellow in Zoology in the University of Illinois. 



1916 




. ... Summer at Marine Biological Laboratory at V'oods Hole, 






Mass. and Harpswell Laboratory, South Harpswell, Me. 


1916 




. ... Elected to the Illinois Chapter of the Society of Sig- 






ma Xi and to the American Microscopical Society. 


1916- 


■1917... Honorary Fellow in Zoology, University of Illinois. 


Publications: 




U ) 


On the Systematic Position of Haplobothrium globulif orme 






PnnnPT* TVftnc Pnv Qrm Pan Qomiqc TTT VTTT CIof»f "1 * 
\j U up ex . 11 all d . Iwjr • uUti . \j dil ., OCT jl c o ill, V 1 1 1 , 06CU.X* 






A — U. J.£7JLffc. 




(2) 


A Mow HracTrtflci "P t» Am fl mi o n o 1 wo T Tttdnc T?rtT7 fon Two^ 
ti HCW UUUc 1 I ULi ixUlXcL LitxXya, Jj . iiclllb. flU.y . \jaHm lllblf 






10(2) :81-119. 3 pis. 1914. 




(3) 


Contributions to the Life-History of Proteocephalus 






ambloplitis. A Parasite of the Black Bass. Contrb. to 






Can. Biol., 47th Rep. Dept. Fisheries, Ottawa, Canada, 






Fasc. II :177-194. 3 pis. 1915. 




(4) 


Trematodes from Marine and Fresh-Water Fishes, includ- 






ing one Species of Ectoparasitic Turbellarian. Trans. 






koy- Soc- Can., Series III, IX :181-2C5, 3 pis* 1915. 




(5) 


Notes on Porocephalus globicephalus . In conjunction 


_ 




with Thesle T. Job. Journ. Parasitology, 3(3) :138.